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<record><TEI><teiHeader><fileDesc><titleStmt><title xml:lang="en">Highlights of the <italic>Didymellaceae</italic>: A polyphasic approach to
characterise <italic>Phoma</italic> and related pleosporalean genera</title><author><name sortKey="Aveskamp, M M" sort="Aveskamp, M M" uniqKey="Aveskamp M" first="M. M." last="Aveskamp">M. M. Aveskamp</name><affiliation><nlm:aff id="aff1"><italic>CBS-KNAW Fungal Biodiversity Centre, Uppsalalaan 8, 3584 CT Utrecht, The Netherlands</italic></nlm:aff></affiliation><affiliation><nlm:aff id="aff3"><italic>Wageningen University and Research Centre (WUR), Laboratory of Phytopathology, Droevendaalsesteeg 1, 6708 PB Wageningen, The Netherlands</italic></nlm:aff></affiliation></author><author><name sortKey="De Gruyter, J" sort="De Gruyter, J" uniqKey="De Gruyter J" first="J." last="De Gruyter">J. De Gruyter</name><affiliation><nlm:aff id="aff1"><italic>CBS-KNAW Fungal Biodiversity Centre, Uppsalalaan 8, 3584 CT Utrecht, The Netherlands</italic></nlm:aff></affiliation><affiliation><nlm:aff id="aff2"><italic>Dutch Plant Protection Service (PD), Geertjesweg 15, 6706 EA Wageningen, The Netherlands</italic></nlm:aff></affiliation></author><author><name sortKey="Woudenberg, J H C" sort="Woudenberg, J H C" uniqKey="Woudenberg J" first="J. H. C." last="Woudenberg">J. H. C. Woudenberg</name><affiliation><nlm:aff id="aff1"><italic>CBS-KNAW Fungal Biodiversity Centre, Uppsalalaan 8, 3584 CT Utrecht, The Netherlands</italic></nlm:aff></affiliation></author><author><name sortKey="Verkley, G J M" sort="Verkley, G J M" uniqKey="Verkley G" first="G. J. M." last="Verkley">G. J. M. Verkley</name><affiliation><nlm:aff id="aff1"><italic>CBS-KNAW Fungal Biodiversity Centre, Uppsalalaan 8, 3584 CT Utrecht, The Netherlands</italic></nlm:aff></affiliation></author><author><name sortKey="Crous, P W" sort="Crous, P W" uniqKey="Crous P" first="P. W." last="Crous">P. W. Crous</name><affiliation><nlm:aff id="aff1"><italic>CBS-KNAW Fungal Biodiversity Centre, Uppsalalaan 8, 3584 CT Utrecht, The Netherlands</italic></nlm:aff></affiliation><affiliation><nlm:aff id="aff3"><italic>Wageningen University and Research Centre (WUR), Laboratory of Phytopathology, Droevendaalsesteeg 1, 6708 PB Wageningen, The Netherlands</italic></nlm:aff></affiliation></author></titleStmt><publicationStmt><idno type="wicri:source">PMC</idno><idno type="pmid">20502538</idno><idno type="pmc">2836210</idno><idno type="url">http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2836210</idno><idno type="RBID">PMC:2836210</idno><idno type="doi">10.3114/sim.2010.65.01</idno><date when="2010">2010</date><idno type="wicri:Area/Pmc/Corpus">000262</idno><idno type="wicri:explorRef" wicri:stream="Pmc" wicri:step="Corpus" wicri:corpus="PMC">000262</idno></publicationStmt><sourceDesc><biblStruct><analytic><title xml:lang="en" level="a" type="main">Highlights of the <italic>Didymellaceae</italic>: A polyphasic approach to
characterise <italic>Phoma</italic> and related pleosporalean genera</title><author><name sortKey="Aveskamp, M M" sort="Aveskamp, M M" uniqKey="Aveskamp M" first="M. M." last="Aveskamp">M. M. Aveskamp</name><affiliation><nlm:aff id="aff1"><italic>CBS-KNAW Fungal Biodiversity Centre, Uppsalalaan 8, 3584 CT Utrecht, The Netherlands</italic></nlm:aff></affiliation><affiliation><nlm:aff id="aff3"><italic>Wageningen University and Research Centre (WUR), Laboratory of Phytopathology, Droevendaalsesteeg 1, 6708 PB Wageningen, The Netherlands</italic></nlm:aff></affiliation></author><author><name sortKey="De Gruyter, J" sort="De Gruyter, J" uniqKey="De Gruyter J" first="J." last="De Gruyter">J. De Gruyter</name><affiliation><nlm:aff id="aff1"><italic>CBS-KNAW Fungal Biodiversity Centre, Uppsalalaan 8, 3584 CT Utrecht, The Netherlands</italic></nlm:aff></affiliation><affiliation><nlm:aff id="aff2"><italic>Dutch Plant Protection Service (PD), Geertjesweg 15, 6706 EA Wageningen, The Netherlands</italic></nlm:aff></affiliation></author><author><name sortKey="Woudenberg, J H C" sort="Woudenberg, J H C" uniqKey="Woudenberg J" first="J. H. C." last="Woudenberg">J. H. C. Woudenberg</name><affiliation><nlm:aff id="aff1"><italic>CBS-KNAW Fungal Biodiversity Centre, Uppsalalaan 8, 3584 CT Utrecht, The Netherlands</italic></nlm:aff></affiliation></author><author><name sortKey="Verkley, G J M" sort="Verkley, G J M" uniqKey="Verkley G" first="G. J. M." last="Verkley">G. J. M. Verkley</name><affiliation><nlm:aff id="aff1"><italic>CBS-KNAW Fungal Biodiversity Centre, Uppsalalaan 8, 3584 CT Utrecht, The Netherlands</italic></nlm:aff></affiliation></author><author><name sortKey="Crous, P W" sort="Crous, P W" uniqKey="Crous P" first="P. W." last="Crous">P. W. Crous</name><affiliation><nlm:aff id="aff1"><italic>CBS-KNAW Fungal Biodiversity Centre, Uppsalalaan 8, 3584 CT Utrecht, The Netherlands</italic></nlm:aff></affiliation><affiliation><nlm:aff id="aff3"><italic>Wageningen University and Research Centre (WUR), Laboratory of Phytopathology, Droevendaalsesteeg 1, 6708 PB Wageningen, 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="2010">2010</date></imprint></series></biblStruct></sourceDesc></fileDesc><profileDesc><textClass></textClass></profileDesc></teiHeader><front><div type="abstract" xml:lang="en"><p>Fungal taxonomists routinely encounter problems when dealing with asexual
fungal species due to poly- and paraphyletic generic phylogenies, and unclear
species boundaries. These problems are aptly illustrated in the genus
<italic>Phoma</italic>. This phytopathologically significant fungal genus is currently
subdivided into nine sections which are mainly based on a single or just a few
morphological characters. However, this subdivision is ambiguous as several of
the section-specific characters can occur within a single species. In
addition, many teleomorph genera have been linked to <italic>Phoma</italic>, three of
which are recognised here. In this study it is attempted to delineate generic
boundaries, and to come to a generic circumscription which is more correct
from an evolutionary point of view by means of multilocus sequence typing.
Therefore, multiple analyses were conducted utilising sequences obtained from
28S nrDNA (Large Subunit - LSU), 18S nrDNA (Small Subunit - SSU), the Internal
Transcribed Spacer regions 1 & 2 and 5.8S nrDNA (ITS), and part of the
β-tubulin (TUB) gene region. A total of 324 strains were included in the
analyses of which most belonged to <italic>Phoma</italic> taxa, whilst 54 to related
pleosporalean fungi. In total, 206 taxa were investigated, of which 159 are
known to have affinities to <italic>Phoma</italic>. The phylogenetic analysis revealed
that the current Boeremaean subdivision is incorrect from an evolutionary
point of view, revealing the genus to be highly polyphyletic. <italic>Phoma</italic>species are retrieved in six distinct clades within the <italic>Pleosporales</italic>,
and appear to reside in different families. The majority of the species,
however, including the generic type, clustered in a recently established
family, <italic>Didymellaceae</italic>. In the second part of this study, the
phylogenetic variation of the species and varieties in this clade was further
assessed. Next to the genus <italic>Didymella</italic>, which is considered to be the
sole teleomorph of <italic>Phoma s. str</italic>., we also retrieved taxa belonging to
the teleomorph genera <italic>Leptosphaerulina</italic> and <italic>Macroventuria</italic> in
this clade. Based on the sequence data obtained, the <italic>Didymellaceae</italic>segregate into at least 18 distinct clusters, of which many can be associated
with several specific taxonomic characters. Four of these clusters were
defined well enough by means of phylogeny and morphology, so that the
associated taxa could be transferred to separate genera. Aditionally, this
study addresses the taxonomic description of eight species and two varieties
that are novel to science, and the recombination of 61 additional taxa.</p></div></front><back><div1 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article-type="research-article"><pmc-dir>properties open_access</pmc-dir>
<front><journal-meta><journal-id journal-id-type="nlm-ta">Stud Mycol</journal-id><journal-id journal-id-type="publisher-id">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">20502538</article-id><article-id pub-id-type="pmc">2836210</article-id><article-id pub-id-type="publisher-id">0001</article-id><article-id pub-id-type="doi">10.3114/sim.2010.65.01</article-id><article-categories><subj-group subj-group-type="heading"><subject>Articles</subject></subj-group></article-categories><title-group><article-title>Highlights of the <italic>Didymellaceae</italic>: A polyphasic approach to
characterise <italic>Phoma</italic> and related pleosporalean genera</article-title></title-group><contrib-group><contrib contrib-type="author"><name><surname>Aveskamp</surname><given-names>M.M.</given-names></name><xref ref-type="aff" rid="aff1">1</xref><xref ref-type="aff" rid="aff3">3</xref><xref ref-type="corresp" rid="cor1">*</xref><xref ref-type="fn" rid="fn1">#</xref></contrib><contrib contrib-type="author"><name><surname>de Gruyter</surname><given-names>J.</given-names></name><xref ref-type="aff" rid="aff1">1</xref><xref ref-type="aff" rid="aff2">2</xref></contrib><contrib contrib-type="author"><name><surname>Woudenberg</surname><given-names>J.H.C.</given-names></name><xref ref-type="aff" rid="aff1">1</xref></contrib><contrib contrib-type="author"><name><surname>Verkley</surname><given-names>G.J.M.</given-names></name><xref ref-type="aff" rid="aff1">1</xref></contrib><contrib contrib-type="author"><name><surname>Crous</surname><given-names>P.W.</given-names></name><xref ref-type="aff" rid="aff1">1</xref><xref ref-type="aff" rid="aff3">3</xref></contrib></contrib-group><aff id="aff1"><label>1</label><italic>CBS-KNAW Fungal Biodiversity Centre, Uppsalalaan 8, 3584 CT Utrecht, The Netherlands</italic></aff><aff id="aff2"><label>2</label><italic>Dutch Plant Protection Service (PD), Geertjesweg 15, 6706 EA Wageningen, The Netherlands</italic></aff><aff id="aff3"><label>3</label><italic>Wageningen University and Research Centre (WUR), Laboratory of Phytopathology, Droevendaalsesteeg 1, 6708 PB Wageningen, The Netherlands</italic></aff><author-notes><corresp id="cor1"><label>*</label><italic>Correspondence</italic>: Maikel M. Aveskamp,
<email>m.aveskamp@cbs.knaw.nl</email></corresp><fn id="fn1"><label>#</label><p><italic>Current address</italic>: Mycolim BV, Veld Oostenrijk 13, 5961 NV Horst,
The Netherlands</p></fn></author-notes><pub-date pub-type="ppub"><year>2010</year></pub-date><volume>65</volume><issue-title>Highlights of the <italic>Didymellaceae</italic>: A polyphasic approach to
characterise <italic>Phoma</italic> and related pleosporalean genera</issue-title><fpage>1</fpage><lpage>60</lpage><permissions><copyright-statement>Copyright © Copyright 2010 CBS Fungal Biodiversity Centre</copyright-statement><copyright-year>2010</copyright-year><license><license-p>You are free to share - to copy, distribute and transmit the work, under
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<ext-link ext-link-type="uri" xlink:href="http://creativecommons.org/licenses/by-nc-nd/3.0/legalcode">http://creativecommons.org/licenses/by-nc-nd/3.0/legalcode.</ext-link> Any of the above
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Nothing in this license impairs or restricts the author's moral rights.</license-p></license></permissions><self-uri xlink:title="pdf" xlink:href="1.pdf"></self-uri><abstract><p>Fungal taxonomists routinely encounter problems when dealing with asexual
fungal species due to poly- and paraphyletic generic phylogenies, and unclear
species boundaries. These problems are aptly illustrated in the genus
<italic>Phoma</italic>. This phytopathologically significant fungal genus is currently
subdivided into nine sections which are mainly based on a single or just a few
morphological characters. However, this subdivision is ambiguous as several of
the section-specific characters can occur within a single species. In
addition, many teleomorph genera have been linked to <italic>Phoma</italic>, three of
which are recognised here. In this study it is attempted to delineate generic
boundaries, and to come to a generic circumscription which is more correct
from an evolutionary point of view by means of multilocus sequence typing.
Therefore, multiple analyses were conducted utilising sequences obtained from
28S nrDNA (Large Subunit - LSU), 18S nrDNA (Small Subunit - SSU), the Internal
Transcribed Spacer regions 1 & 2 and 5.8S nrDNA (ITS), and part of the
β-tubulin (TUB) gene region. A total of 324 strains were included in the
analyses of which most belonged to <italic>Phoma</italic> taxa, whilst 54 to related
pleosporalean fungi. In total, 206 taxa were investigated, of which 159 are
known to have affinities to <italic>Phoma</italic>. The phylogenetic analysis revealed
that the current Boeremaean subdivision is incorrect from an evolutionary
point of view, revealing the genus to be highly polyphyletic. <italic>Phoma</italic>species are retrieved in six distinct clades within the <italic>Pleosporales</italic>,
and appear to reside in different families. The majority of the species,
however, including the generic type, clustered in a recently established
family, <italic>Didymellaceae</italic>. In the second part of this study, the
phylogenetic variation of the species and varieties in this clade was further
assessed. Next to the genus <italic>Didymella</italic>, which is considered to be the
sole teleomorph of <italic>Phoma s. str</italic>., we also retrieved taxa belonging to
the teleomorph genera <italic>Leptosphaerulina</italic> and <italic>Macroventuria</italic> in
this clade. Based on the sequence data obtained, the <italic>Didymellaceae</italic>segregate into at least 18 distinct clusters, of which many can be associated
with several specific taxonomic characters. Four of these clusters were
defined well enough by means of phylogeny and morphology, so that the
associated taxa could be transferred to separate genera. Aditionally, this
study addresses the taxonomic description of eight species and two varieties
that are novel to science, and the recombination of 61 additional taxa.</p></abstract><kwd-group><kwd><italic>Boeremia</italic></kwd><kwd>coelomycetes</kwd><kwd><italic>Didymella</italic></kwd><kwd><italic>Didymellaceae</italic></kwd><kwd>DNA phylogeny</kwd><kwd><italic>Epicoccum</italic></kwd><kwd><italic>Leptosphaerulina</italic></kwd><kwd><italic>Macroventuria</italic></kwd><kwd><italic>Peyronellaea</italic></kwd><kwd><italic>Phoma</italic></kwd><kwd><italic>Pleosporales</italic></kwd><kwd>taxonomy</kwd><kwd><italic>Stagonosporopsis</italic></kwd></kwd-group></article-meta><notes><fn-group><fn><p><bold>Taxonomic novelties: New genus:</bold> <italic>Boeremia</italic> Aveskamp, Gruyter
& Verkley. <bold>New species:</bold> <italic>Phoma brasiliensis</italic> Aveskamp,
Gruyter & Verkley, <italic>Ph. bulgarica</italic> Aveskamp, Gruyter & Verkley,
<italic>Ph. dactylidis</italic> Aveskamp, Gruyter & Verkley, <italic>Ph. dimorpha</italic>Aveskamp, Gruyter & Verkley, <italic>Ph. longicolla</italic> Aveskamp, Gruyter
& Verkley, <italic>Ph. minor</italic> Aveskamp, Gruyter & Verkley, <italic>Ph.
pedeiae</italic> Aveskamp, Gruyter & Verkley, <italic>Ph. saxea</italic> Aveskamp,
Gruyter & Verkley. <bold>New varieties:</bold> <italic>Boeremia exigua</italic> var.
<italic>gilvescens</italic> Aveskamp, Gruyter & Verkley, <italic>B. exigua</italic> var.
<italic>pseudolilacis</italic> Aveskamp, Gruyter & Verkley. <bold>New
combinations:</bold> <italic><bold>Boeremia</bold> crinicola</italic> (Siemasko) Aveskamp,
Gruyter & Verkley, <italic>B. diversispora</italic> (Bubák) Aveskamp,
Gruyter & Verkley, <italic>B. exigua</italic> var. <italic>exigua</italic> (Desm.)
Aveskamp, Gruyter & Verkley, <italic>B. exigua</italic> var. <italic>heteromorpha</italic>(Schulzer & Sacc.) Aveskamp, Gruyter & Verkley, <italic>B. exigua</italic>var. <italic>lilacis</italic> (Sacc.) Aveskamp, Gruyter & Verkley, <italic>B.
exigua</italic> var. <italic>linicola</italic> (Naumov & Vassiljevsky) Aveskamp,
Gruyter & Verkley, <italic>B. exigua</italic> var. <italic>populi</italic> (Gruyter &
Scheer) Aveskamp, Gruyter & Verkley, <italic>B. exigua</italic> var.
<italic>coffeae</italic> (Henn.) Aveskamp, Gruyter & Verkley, <italic>B. exigua</italic>var. <italic>viburni</italic> (Roum. ex. Sacc.) Aveskamp, Gruyter & Verkley,
<italic>B. foveata</italic> (Foister) Aveskamp, Gruyter & Verkley, <italic>B.
lycopersici</italic> (Cooke) Aveskamp, Gruyter & Verkley, <italic>B.
noackiana</italic> (Allesch.) Aveskamp, Gruyter & Verkley, <italic>B.
sambuci-nigrae</italic> (Sacc.) Aveskamp, Gruyter & Verkley, <italic>B.
strasseri</italic> (Moesz) Aveskamp, Gruyter & Verkley, <italic>B. telephii</italic>(Vestergr.) Aveskamp, Gruyter & Verkley, <italic><bold>Epicoccum</bold>pimprinum</italic> (P.N. Mathur, S.K. Menon & Thirum.) Aveskamp, Gruyter &
Verkley, <italic>E. sorghi</italic> (Sacc.) Aveskamp, Gruyter & Verkley,
<italic><bold>Peyronellaea</bold> americana</italic> (Morgan-Jones & J.F. White)
Aveskamp, Gruyter & Verkley, <italic>Pey. alectorolophi</italic> (Rehm.) Aveskamp,
Gruyter & Verkley, <italic>Pey. anserina</italic> (Marchal) Aveskamp, Gruyter
& Verkley, <italic>Pey. arachidicola</italic> (Khokhr.) Aveskamp, Gruyter &
Verkley, <italic>Pey. aurea</italic> (Gruyter, Noordel. & Boerema) Aveskamp,
Gruyter & Verkley, <italic>Pey. calorpreferens</italic> (Boerema, Gruyter &
Noordel.) Aveskamp, Gruyter & Verkley, <italic>Pey. coffeae-arabicae</italic>(Aveskamp, Verkley & Gruyter) Aveskamp, Gruyter & Verkley, <italic>Pey.
curtisii</italic> (Berk.) Aveskamp, Gruyter & Verkley, <italic>Pey.
eucalyptica</italic> (Sacc.) Aveskamp, Gruyter & Verkley, <italic>Pey.
gardeniae</italic> (S. Chandra & Tandon) Aveskamp, Gruyter & Verkley,
<italic>Pey. lethalis</italic> (Ellis & Bartholomew) Aveskamp, Gruyter &
Verkley, <italic>Pey. pomorum</italic> var. <italic>pomorum</italic> (Thüm.) Aveskamp,
Gruyter & Verkley, <italic>Pey. pomorum</italic> var. <italic>circinata</italic>(Kusnezowa) Aveskamp, Gruyter & Verkley, <italic>Pey. pomorum</italic> var.
<italic>cyanea</italic> (Jooste & Papendorf) Aveskamp, Gruyter & Verkley,
<italic>Pey. obtusa</italic> (Fuckel) Aveskamp, Gruyter & Verkley, <italic>Pey.
pinodella</italic> (L.K. Jones) Aveskamp, Gruyter & Verkley, <italic>Pey.
pinodes</italic> (Berk. & A. Bloxam) Aveskamp, Gruyter & Verkley, <italic>Pey.
protuberans</italic> (Lév.) Aveskamp, Gruyter & Verkley, <italic>Pey.
sancta</italic> (Aveskamp, Gruyter & Verkley) Aveskamp, Gruyter & Verkley,
<italic>Pey. subglomerata</italic> (Boerema, Gruyter & Noordel.) Aveskamp, Gruyter
& Verkley, <italic>Pey. zeae-maydis</italic> (Arny & R.R. Nelson)Aveskamp,
Gruyter & Verkley, <italic><bold>Phoma</bold> clematidis-rectae</italic> (Petr.)
Aveskamp, Woudenberg & Gruyter, <italic>Ph. noackiana</italic> (Allesch.)
Aveskamp, Gruyter & Verkley, <italic><bold>Stagonosporopsis</bold> ajacis</italic>(Thüm.) Aveskamp, Gruyter & Verkley, <italic>S. andigena</italic> (Turkenst.)
Aveskamp, Gruyter & Verkley, <italic>S. artemisiicola</italic> (Hollós)
Aveskamp, Gruyter & Verkley, <italic>S. astragali</italic> (Cooke & Harkn.)
Aveskamp, Gruyter & Verkley, <italic>S. caricae</italic> (Sydow & P. Sydow)
Aveskamp, Gruyter & Verkley, <italic>S. crystalliniformis</italic> (Loer., R.
Navarro, M. Lôbo & Turkenst.) Aveskamp, Gruyter & Verkley,
<italic>S. cucurbitacearum</italic> (Fr.) Aveskamp, Gruyter & Verkley, <italic>S.
dorenboschii</italic> (Noordel. & Gruyter) Aveskamp, Gruyter & Verkley,
<italic>S. heliopsidis</italic> (H.C. Greene) Aveskamp, Gruyter & Verkley, <italic>S.
ligulicola</italic> var. <italic>ligulicola</italic> (K.F. Baker, Dimock & L.H. Davis)
Aveskamp, Gruyter & Verkley, <italic>S. ligulicola</italic> var.
<italic>inoxydabilis</italic> (Boerema) Aveskamp, Gruyter & Verkley, <italic>S.
loticola</italic> (Died.) Aveskamp, Gruyter & Verkley, <italic>S.
oculo-hominis</italic> (Punith.) Aveskamp, Gruyter & Verkley, <italic>S.
rudbeckiae</italic> (Fairm.) Aveskamp, Gruyter & Verkley, <italic>S.
trachelii</italic> (Allesch.) Aveskamp, Gruyter & Verkley, <italic>S.
valerianellaea</italic> (Gindrat, Semecnik & Bolay) Aveskamp, Gruyter &
Verkley. <bold>New names:</bold> <italic><bold>Peyronellaea</bold> australis</italic> Aveskamp,
Gruyter & Verkley, <italic><bold>Phoma</bold> fungicola</italic> Aveskamp, Gruyter &
Verkley, <italic>Ph. novae-verbascicola</italic> Aveskamp, Gruyter & Verkley.</p></fn></fn-group></notes></front><body><sec><title>INTRODUCTION</title><p>Coelomycetous fungi (<xref ref-type="bibr" rid="ref93">Grove
1935</xref>) are geographically widespread and are found in numerous
ecological niches. Sutton
(<xref ref-type="bibr" rid="ref219">1980</xref>) mentions exponents of
this anamorph group inhabiting soil, organic debris, and water, as well as
species that parasitise other fungi, lichens, insects and vertebrates. A
substantial percentage of the coelomycetes is associated with plant material,
either as opportunists or as primary pathogens
(<xref ref-type="bibr" rid="ref219">Sutton 1980</xref>).</p><p>Difficulties in morphological identification have resulted in a poor
understanding of the generic and species boundaries in the coelomycetes
(Sutton <xref ref-type="bibr" rid="ref218">1977</xref>,
<xref ref-type="bibr" rid="ref219">1980</xref>,
<xref ref-type="bibr" rid="ref146">Nag Raj 1981</xref>,
<xref ref-type="bibr" rid="ref6">Van der Aa <italic>et al.</italic>1990</xref>, Torres <italic>et al.</italic><xref ref-type="bibr" rid="ref225">2005a</xref>,
<xref ref-type="bibr" rid="ref226">b</xref>,
<xref ref-type="bibr" rid="ref100">De Gruyter <italic>et al</italic>.
2009</xref>). In an attempt to improve the classification of the
coelomycetes, Sutton (<xref ref-type="bibr" rid="ref219">1980</xref>)
proposed to divide the order into six suborders, which unfortunately proved to
be highly artificial from an evolutionary perspective
(<xref ref-type="bibr" rid="ref100">De Gruyter <italic>et al</italic>.
2009</xref>).</p><p>The current common procedure for isolate identification, which chiefly
relies on similarity of DNA sequences to those found in public DNA libraries
(<xref ref-type="bibr" rid="ref115">Hyde & Soytong 2007</xref>),
combined with the high level of incorrectly identified sequences in these
databases (Bridge <italic>et al.</italic><xref ref-type="bibr" rid="ref59">2003</xref>,
<xref ref-type="bibr" rid="ref60">2004</xref>,
<xref ref-type="bibr" rid="ref150">Nilsson <italic>et al.</italic> 2006</xref>)
placed the likelihood of achieving correct identifications of coelomycetous
fungi under intense scrutiny. As pointed out by De Gruyter <italic>et al.</italic>(<xref ref-type="bibr" rid="ref100">2009</xref>), for appropriate
morphological identifications within the coelomycete genera <italic>in vitro</italic>studies are essential, for example in the cases in which quarantine pathogens
are involved (<xref ref-type="bibr" rid="ref18">Aveskamp <italic>et al.</italic>2008</xref>). For the current generic delimitation of this class, the use
of conidiogenesis characters as taxonomic criteria is of major importance
(<xref ref-type="bibr" rid="ref114">Hughes 1953</xref>;
<xref ref-type="bibr" rid="ref30">Boerema 1965</xref>,
<xref ref-type="bibr" rid="ref36">Boerema & Bollen 1975</xref>,
Sutton <xref ref-type="bibr" rid="ref217">1964</xref>,
<xref ref-type="bibr" rid="ref218">1977</xref>,
<xref ref-type="bibr" rid="ref219">1980</xref>,
<xref ref-type="bibr" rid="ref206">Singh <italic>et al.</italic>1997</xref>).</p><sec><title>Phoma</title><p>The genus <italic>Phoma</italic> Sacc. emend. Boerema & G.J. Bollen
(<italic>Pleosporales</italic>) is a good example of a coelomycetous genus made
fascinating by its great ecological diversity, but taxing investigators with
profound difficulties in making identifications. The majority of the taxa
within this mitosporic genus have been found in association with land plants,
causing mainly leaf and stem spots
(<xref ref-type="bibr" rid="ref18">Aveskamp <italic>et al</italic>.
2008</xref>, <xref ref-type="bibr" rid="ref242">Zhang <italic>et al.</italic>2009</xref>). Approximately 50 % of the <italic>Phoma</italic> taxa that were
redescribed by Boerema <italic>et al.</italic>(<xref ref-type="bibr" rid="ref55">2004</xref>) are recognised as
relevant phytopathogenic fungi, including a series of pathogens with
quarantine status (<xref ref-type="bibr" rid="ref55">Boerema <italic>et al.</italic>2004</xref>, <xref ref-type="bibr" rid="ref18">Aveskamp <italic>et al</italic>.
2008</xref>). Although most taxa are continuously present in the
environment as saprobic soil organisms, many species switch to a pathogenic
lifestyle when a suitable host is encountered
(<xref ref-type="bibr" rid="ref18">Aveskamp <italic>et al</italic>.
2008</xref>). The genus further comprises several species and varieties
that are recognised as endophytic, fungicolous and lichenicolous fungi
(<italic>e.g.</italic> <xref ref-type="bibr" rid="ref103">Hawksworth
1981</xref>, <xref ref-type="bibr" rid="ref240">Xianshu <italic>et al</italic>.
1994</xref>, <xref ref-type="bibr" rid="ref214">Sullivan & White
2000</xref>, <xref ref-type="bibr" rid="ref104">Hawksworth & Cole
2004</xref>, <xref ref-type="bibr" rid="ref81">Diederich <italic>et
al.</italic> 2007</xref>, <xref ref-type="bibr" rid="ref194">Schoch <italic>et
al</italic>. 2009a</xref>). In addition, approximately 10 species are known
as pathogens of humans (<italic>e.g.</italic> <xref ref-type="bibr" rid="ref108">De
Hoog <italic>et al</italic>. 2000</xref>,
<xref ref-type="bibr" rid="ref21">Balis <italic>et al.</italic> 2006</xref>)
and other vertebrates, such as cattle
(<xref ref-type="bibr" rid="ref68">Costa <italic>et al.</italic> 1993</xref>)
and fish (<xref ref-type="bibr" rid="ref184">Ross <italic>et al</italic>.
1975</xref>, <xref ref-type="bibr" rid="ref102">Hatai <italic>et al</italic>.
1986</xref>, <xref ref-type="bibr" rid="ref232">Voronin
1989</xref>, <xref ref-type="bibr" rid="ref86">Faisal <italic>et al</italic>.
2007</xref>). Next to such an active role in vertebrate pathology,
<italic>Phoma</italic> spp. may indirectly affect animal health by the production of
toxic secondary metabolites (<xref ref-type="bibr" rid="ref26">Bennett
1983</xref>, <xref ref-type="bibr" rid="ref162">Pedras & Biesenthal
2000</xref>, <xref ref-type="bibr" rid="ref176">Rai <italic>et al.</italic>2009</xref>), as is known for <italic>Ph. sorghina</italic> in straw roofs in
South Africa (<xref ref-type="bibr" rid="ref174">Rabie <italic>et al.</italic>1975</xref>) and may be the case in <italic>Ph. pomorum</italic> in cattle feed
(<xref ref-type="bibr" rid="ref209">Sørensen <italic>et al.</italic>2009</xref>). An almost completely unexplored habitat of <italic>Phoma</italic>spp. is the marine environment (<xref ref-type="bibr" rid="ref128">Kohlmeyer
& Volkmann-Kohlmeyer 1991</xref>), in which <italic>Phoma</italic> species
are regularly found that are completely new to science (<italic>e.g.</italic><xref ref-type="bibr" rid="ref156">Osterhage <italic>et al</italic>.
2000</xref>, <xref ref-type="bibr" rid="ref241">Yarden <italic>et al</italic>.
2007</xref>).</p><p>The genus <italic>Phoma</italic> has always been considered to be one of the
largest fungal genera, with more than 3 000 infrageneric taxa described
(<xref ref-type="bibr" rid="ref137">Monte <italic>et al</italic>. 1991</xref>).
The number of species described in <italic>Phoma</italic> rose to this level due to
the common practice of host associated nomenclature, in combination with the
paucity in micromorphological characters and a high variability in cultural
characteristics. These factors have resulted in the fact that the systematics
of the genus never has been fully understood
(<xref ref-type="bibr" rid="ref18">Aveskamp <italic>et al.</italic>2008</xref>). Based on various morphological features depicted by earlier
workers, probably less than one-tenth of the 3 200 species listed in MycoBank
(<ext-link ext-link-type="uri" xlink:href="www.mycobank.org">www.mycobank.org</ext-link>,
<xref ref-type="bibr" rid="ref72">Crous <italic>et al.</italic> 2004</xref>,
<xref ref-type="bibr" rid="ref182">Robert <italic>et al.</italic> 2005</xref>)
can currently still be recognised as a separate <italic>Phoma</italic> taxon. Many of
those names were thus already reduced to synonymy after an extensive study of
the genus (<xref ref-type="bibr" rid="ref55">Boerema <italic>et al</italic>.
2004</xref>), and after a thoroughly revised generic concept of the
morphologically similar genera <italic>Ascochyta</italic>(<xref ref-type="bibr" rid="ref36">Boerema & Bollen 1975</xref>)
and <italic>Phyllosticta</italic> (<xref ref-type="bibr" rid="ref2">Van der Aa
1973</xref>, <xref ref-type="bibr" rid="ref5">Van der Aa & Vanev
2002</xref>). Many other species could be recombined into other
coelomycete genera, such as <italic>Asteromella, Microsphaeropsis, Phomopsis,
Pleurophoma, Pyrenochaeta</italic> and <italic>Stagonospora</italic> (Sutton
<xref ref-type="bibr" rid="ref217">1964</xref>,
<xref ref-type="bibr" rid="ref219">1980</xref>,
<xref ref-type="bibr" rid="ref36">Boerema & Bollen 1975</xref>). In
addition, <italic>Coniothyrium</italic> and <italic>Paraconiothyrium</italic> have regularly
been mistaken for <italic>Phoma</italic> (<xref ref-type="bibr" rid="ref228">Verkley
<italic>et al.</italic> 2004</xref>, <xref ref-type="bibr" rid="ref78">Damm
<italic>et al.</italic> 2008</xref>,
<xref ref-type="bibr" rid="ref239">Woudenberg <italic>et al</italic>.
2009</xref>). In their studies, Boerema <italic>et al.</italic>(<xref ref-type="bibr" rid="ref55">2004</xref>) recognised a total of
215 <italic>Phoma</italic> taxa and eight teleomorph species with an unnamed
<italic>Phoma</italic> anamorph, although this is probably just the tip of the iceberg
as, thus far, only 40 % of the herbarium species mentioned in literature could
be recovered and studied properly. Additionally, novel species are described
regularly in this genus (<italic>e.g.</italic><xref ref-type="bibr" rid="ref104">Hawksworth & Cole 2004</xref>,
<xref ref-type="bibr" rid="ref225">Torres <italic>et al.</italic> 2005a</xref>,
<xref ref-type="bibr" rid="ref131">Li <italic>et al.</italic> 2006</xref>,
<xref ref-type="bibr" rid="ref81">Diederich <italic>et al.</italic>2007</xref>, <xref ref-type="bibr" rid="ref19">Aveskamp <italic>et al.</italic>2009a</xref>, <xref ref-type="bibr" rid="ref80">Davidson <italic>et
al.</italic> 2009</xref>).</p><p>A subdivision of the asexual genus <italic>Phoma</italic> that is currently widely
applied divides the genus into nine sections, including the sections
<italic>Phoma, Heterospora, Macrospora, Paraphoma, Peyronellaea, Phyllostictoides,
Pilosa, Plenodomus</italic> and <italic>Sclerophomella</italic>(<xref ref-type="bibr" rid="ref34">Boerema 1997</xref>). These sections
are primarily based on just a few morphological or physiological characters
and have not been confirmed as biologically realistic by molecular biological
studies. The number of taxa per section may vary, ranging from almost 70
species in section <italic>Phoma</italic> to only two in section <italic>Pilosa</italic>. In
<xref ref-type="table" rid="tbl1">Table 1</xref>, a list is provided
with the main characters of every section
(<xref ref-type="bibr" rid="ref34">Boerema 1997</xref>). This
subdivision into sections has led to an identification system that is
considered to be extremely helpful in morphological identification
(<xref ref-type="bibr" rid="ref55">Boerema <italic>et al.</italic>2004</xref>). However, as was hypothesised by Boerema <italic>et al.</italic>(<xref ref-type="bibr" rid="ref55">2004</xref>), the classification has
proved to be artificial. Molecular evidence has shown that the sections are
linked to phylogenetically distinct teleomorph genera
(<xref ref-type="bibr" rid="ref178">Reddy <italic>et al</italic>. 1998</xref>,
<xref ref-type="bibr" rid="ref226">Torres <italic>et al.</italic> 2005b</xref>,
<xref ref-type="bibr" rid="ref100">De Gruyter <italic>et al</italic>.
2009</xref>). Even these teleomorph genera are not always monophyletic
(<xref ref-type="bibr" rid="ref139">Morales <italic>et al.</italic>1995</xref>, <xref ref-type="bibr" rid="ref62">Câmara <italic>et
al.</italic> 2002</xref>, <xref ref-type="bibr" rid="ref126">Kodsueb <italic>et
al</italic>. 2006</xref>, <xref ref-type="bibr" rid="ref116">Inderbitzin
<italic>et al.</italic> 2009</xref>). In addition, characters that are thought to
be specific for a certain section appeared to be polyphyletic, as is
illustrated for dictyochlamydospores and setose pycnidia, the main characters
for the sections <italic>Peyronellaea</italic>(<xref ref-type="bibr" rid="ref19">Aveskamp <italic>et al.</italic>2009a</xref>) and <italic>Paraphoma</italic>(<xref ref-type="bibr" rid="ref92">Grondona <italic>et al</italic>.
1997</xref>, <xref ref-type="bibr" rid="ref101">De Gruyter <italic>et
al.</italic> 2010</xref>) respectively. Furthermore, <italic>Phoma</italic> section
<italic>Phoma</italic>, a group of species which is characterised by the absence of
chlamydospores, septate conidia, and pycnidial ornamentation or wall
thickening, is considered to be a repository for degenerated and
insufficiently understood species that could not be placed elsewhere.</p><p><table-wrap position="float" id="tbl1"><label>Table 1.</label><caption><p>Overview of the characters of the various <italic>Phoma</italic> sections in the
Boeremaean classification system. Adapted from Boerema <italic>et al.</italic>(<xref ref-type="bibr" rid="ref55">2004</xref>).</p></caption><table frame="hsides" rules="groups"><thead><tr><th valign="top" align="left"><bold>Section</bold></th><th valign="top" align="center"><bold>Teleomorph</bold></th><th valign="top" align="center"><bold>Synanamorph</bold></th><th valign="top" align="center"><bold>Sectional character</bold></th></tr></thead><tbody><tr><td align="left" valign="top"><italic>Heterospora</italic></td><td align="left" valign="top"> —
</td><td align="left" valign="top"><italic>Stagonosporopsis</italic></td><td align="left" valign="top"> Production of distinctly large conidia in addition to the regular conidia
</td></tr><tr><td align="left" valign="top"><italic>Macrospora</italic></td><td align="left" valign="top"><italic>Mycosphaerella</italic></td><td align="left" valign="top"> —
</td><td align="left" valign="top"> Conidia large, measuring 8-19 × 3-7 μm
</td></tr><tr><td align="left" valign="top"><italic>Paraphoma</italic></td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> Setose pycnidia
</td></tr><tr><td align="left" valign="top"><italic>Peyronellaea</italic></td><td align="left" valign="top"> —
</td><td align="left" valign="top"><italic>Epicoccum</italic><xref ref-type="table-fn" rid="tblfn1">*</xref></td><td align="left" valign="top"> Multicellular chlamydospores
</td></tr><tr><td align="left" valign="top"><italic>Phoma</italic></td><td align="left" valign="top"><italic>Didymella</italic></td><td align="left" valign="top"><italic>Phialophora</italic><xref ref-type="table-fn" rid="tblfn1">*</xref></td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"><italic>Phyllostictoides</italic></td><td align="left" valign="top"><italic>Didymella</italic></td><td align="left" valign="top"> —
</td><td align="left" valign="top"> Small septate conidia in addition to the regular conidia
</td></tr><tr><td align="left" valign="top"><italic>Pilosa</italic></td><td align="left" valign="top"><italic>Pleospora</italic></td><td align="left" valign="top"> —
</td><td align="left" valign="top"> Pycnidia covered by pilose outgrows
</td></tr><tr><td align="left" valign="top"><italic>Plenodomus</italic></td><td align="left" valign="top"><italic>Leptosphaeria</italic></td><td align="left" valign="top"><italic>Sclerotium</italic><xref ref-type="table-fn" rid="tblfn1">*</xref></td><td align="left" valign="top"> Pycnidia scleroplectenchymatous
</td></tr><tr><td align="left" valign="top"></td><td align="left" valign="top"></td><td align="left" valign="top"><italic>Phialophora</italic><xref ref-type="table-fn" rid="tblfn1">*</xref></td><td align="left" valign="top"></td></tr><tr><td align="left" valign="top"><italic>Sclerophomella</italic></td><td align="left" valign="top"><italic>Didymella</italic></td><td align="left" valign="top">—
</td><td align="left" valign="top">Pycnidia thick-walled
</td></tr></tbody></table><table-wrap-foot><fn id="tblfn1"><label>*</label><p>Synanamorph only recorded in a single species.</p></fn></table-wrap-foot></table-wrap></p><p>The genus <italic>Phoma</italic> is typified by <italic>Phoma herbarum</italic>(<xref ref-type="bibr" rid="ref29">Boerema 1964</xref>). This species
has thus far not been linked to any teleomorph, but several other species that
are currently accommodated in <italic>Phoma</italic> do have a sexual state. The
species in the section <italic>Pilosa</italic> are linked to the teleomorph genus
<italic>Pleospora</italic>, while many species in the section <italic>Plenodomus</italic> have
a sexual state in <italic>Leptosphaeria</italic>. As mentioned above,
<italic>Leptosphaeria</italic> is para- or possibly polyphyletic
(<xref ref-type="bibr" rid="ref139">Morales <italic>et al.</italic>1995</xref>, <xref ref-type="bibr" rid="ref62">Câmara <italic>et
al.</italic> 2002</xref>). A teleomorph in the poorly studied genus
<italic>Didymella</italic> is associated with approximately 40 <italic>Phoma</italic> species
placed in sections <italic>Phoma, Phyllostictoides</italic> and
<italic>Sclerophomella</italic> (<xref ref-type="bibr" rid="ref55">Boerema <italic>et
al</italic>. 2004</xref>). Moreover, <italic>Phoma</italic> has been linked in
literature to several other teleomorph genera, such as <italic>Mycosphaerella</italic>(<xref ref-type="bibr" rid="ref69">Corlett 1991</xref>,
<xref ref-type="bibr" rid="ref94">De Gruyter 2002</xref>, Crous <italic>et
al</italic>. <xref ref-type="bibr" rid="ref74">2009a</xref>,
<xref ref-type="bibr" rid="ref75">b</xref>), <italic>Belizeana</italic>(<xref ref-type="bibr" rid="ref127">Kohlmeyer & Volkmann-Kohlmeyer
1987</xref>), <italic>Atradidymella</italic>(<xref ref-type="bibr" rid="ref79">Davey & Currah 2009</xref>) and
<italic>Fenestella, Cucurbitaria, Preussia</italic>, and <italic>Westerdykella</italic>(<xref ref-type="bibr" rid="ref14">Von Arx 1981</xref>,
<xref ref-type="bibr" rid="ref242">Zhang <italic>et al</italic>. 2009</xref>).
None of these hypothesised teleomorph-anamorph linkages is supported by
molecular evidence. All must be investigated by study of type material.
However, these associations are unlikely as the mentioned teleomorph genera
are not linked to the <italic>Pleosporales</italic>. The species and teleomorph
relations are also not recognised by Boerema <italic>et al</italic>.
(<xref ref-type="bibr" rid="ref55">2004</xref>), except for two
<italic>Phoma</italic> species of the section <italic>Macrospora, Ph. rabiei</italic> and
<italic>Ph. zeae-maydis</italic> which were linked to
“<italic>Mycosphaerella</italic>” teleomorphs as <italic>M. rabiei</italic>(<xref ref-type="bibr" rid="ref122">Kaiser 1997</xref>,
<xref ref-type="bibr" rid="ref94">De Gruyter 2002</xref>) and <italic>M.
zeae-maydis</italic> (<xref ref-type="bibr" rid="ref142">Mukunya & Boothroid
1973</xref>) respectively. Both species also have names in
<italic>Didymella</italic>. The use of those names is recommended, since
<italic>Mycosphaerella</italic> has been shown to be phylogenetically widely separated
from all known <italic>Phoma</italic> species (<xref ref-type="bibr" rid="ref100">De
Gruyter <italic>et al.</italic> 2009</xref>, Crous <italic>et al</italic>.
<xref ref-type="bibr" rid="ref74">2009a</xref>,
<xref ref-type="bibr" rid="ref75">b</xref>).</p><p>Characteristic strains of the genus concerned have been used in a
Multilocus Sequence Typing (MLST) study of the <italic>Dothideomycetes</italic>, which
indicated that <italic>Phoma</italic> is phylogenetically embedded in the
<italic>Pleosporales</italic> (Schoch <italic>et al</italic>.
<xref ref-type="bibr" rid="ref193">2006</xref>,
<xref ref-type="bibr" rid="ref195">2009b</xref>,
<xref ref-type="bibr" rid="ref242">Zhang <italic>et al</italic>. 2009</xref>).
A similar, but smaller scale study aiming to delineate the species in the
unofficial suborder Phialopycnidiineae
(<xref ref-type="bibr" rid="ref219">Sutton 1980</xref>), revealed that
<italic>Phoma</italic> is highly polyphyletic, as reference species of the various
sections were recovered in distinct clades of the reconstructed phylogeny
(<xref ref-type="bibr" rid="ref100">De Gruyter <italic>et al.</italic>2009</xref>). Type species of the sections <italic>Heterospora, Plenodomus,
Paraphoma</italic> and <italic>Pilosa</italic> appeared to be ancestral to a cluster
comprising types of the other sections, as well as to members of the anamorph
genera <italic>Ascochyta, Microsphaeropsis, Chaetasbolisia, Coniothyrium</italic> and
<italic>Paraconiothyrium.</italic> This group has been elevated to family level and is
now recognised as the <italic>Didymellaceae</italic>(<xref ref-type="bibr" rid="ref100">De Gruyter <italic>et al.</italic>2009</xref>). A B<sc>last</sc>-search in public sequence libraries
revealed a high genetic similarity between species ascribed to the
<italic>Didymellaceae</italic> and two other teleomorph genera, <italic>Macroventuria</italic>and <italic>Leptosphaerulina</italic>, although these genera are morphologically
clearly distinct from <italic>Didymella</italic>(<xref ref-type="bibr" rid="ref1">Van der Aa 1971</xref>,
<xref ref-type="bibr" rid="ref14">Von Arx 1981</xref>,
<xref ref-type="bibr" rid="ref242">Zhang <italic>et al</italic>. 2009</xref>).
The genetic similarity between those two genera has been observed before by
Kodsueb <italic>et al.</italic>(<xref ref-type="bibr" rid="ref126">2006</xref>), but the phylogenetic
relationship with the genus <italic>Didymella</italic> was not noted in their study.
Members of these two genera have therefore also been included in this
study.</p><p>To solve the problems in quarantine species identification of isolates
taken from samples obtained during phytosanitary border controls, a
comprehensive taxonomic system is required
(<xref ref-type="bibr" rid="ref18">Aveskamp <italic>et al.</italic>2008</xref>). As DNA-based techniques do become more and more important
in identification and detection of plant pathogens
(<xref ref-type="bibr" rid="ref58">Bridge 2002</xref>), such a
taxonomic system should be in line with sequence data. One of the major
initiatives in this field is the development of DNA Barcodes
(<xref ref-type="bibr" rid="ref105">Hebert <italic>et al.</italic> 2003</xref>,
<xref ref-type="bibr" rid="ref215">Summerbell <italic>et al</italic>.
2005</xref>), which has been promising in the rapid detection of
potentially serious plant pathogens
(<xref ref-type="bibr" rid="ref17">Armstrong & Ball
2005</xref>).</p><p>Three genes have in recent years been proposed as standard loci for use in
DNA barcoding in fungi. These comprise the internal transcribed spacers (ITS)
of the rDNA operon ITS region (<xref ref-type="bibr" rid="ref84">Druzhinina
<italic>et al.</italic> 2005</xref>), actin (ACT,
<xref ref-type="bibr" rid="ref20">Aveskamp <italic>et al.</italic>2009b</xref>), and cytochrome <italic>c</italic> oxidase subunit I (COI,
<xref ref-type="bibr" rid="ref198">Seifert <italic>et al.</italic>2007</xref>). The last locus was successfully applied in DNA Barcoding of
<italic>Penicillium</italic> (<xref ref-type="bibr" rid="ref198">Seifert <italic>et
al.</italic> 2007</xref>, <xref ref-type="bibr" rid="ref65">Chen <italic>et
al.</italic> 2009</xref>). However, COI analysis applied to a subset of
<italic>Ph. exigua</italic> related strains, did not reveal taxon-specific conserved
SNPs (<xref ref-type="bibr" rid="ref20">Aveskamp <italic>et al.</italic>2009b</xref>), whilst in an attempt to barcode <italic>Aspergillus</italic>, COI
was found to have limited value (<xref ref-type="bibr" rid="ref90">Geiser
<italic>et al.</italic> 2007</xref>). Although ACT has proven helpful in
resolving the phylogeny of <italic>Phoma exigua</italic> below species level
(<xref ref-type="bibr" rid="ref20">Aveskamp <italic>et al.</italic>2009b</xref>), it could not be applied in the present study, as
interspecific variation proved to be too high to align the obtained sequences
properly. The use of ITS as fungal barcode locus is most popular
(<xref ref-type="bibr" rid="ref197">Seifert 2009</xref>) and has been
applied in several taxonomic groups, such as <italic>Trichoderma</italic> and
<italic>Hypocrea</italic> (<xref ref-type="bibr" rid="ref84">Druzhinina <italic>et
al.</italic> 2005</xref>), and <italic>Trichophyton</italic>(<xref ref-type="bibr" rid="ref216">Summerbell <italic>et al.</italic>2007</xref>) and in ecological groups such as wood-inhabiting fungi
(<xref ref-type="bibr" rid="ref147">Naumann <italic>et al.</italic>2007</xref>). The power of this locus for barcoding lies in the multiple
copies that are present within each cell; this phenomenon results in lower
detection thresholds than can be obtained with single-copy loci. Despite the
general practicality of using ITS in barcoding, the locus is relatively
conservative and may oversimplify species delimitations or blur generic
boundaries in some groups (<xref ref-type="bibr" rid="ref151">Nilsson <italic>et
al.</italic> 2008</xref>). In the present study, a combination of four loci
is therefore applied. These include two loci that are renowned for their
capacity to resolve phylogenies above family level, namely parts of the LSU
(Large Subunit – 28S) and SSU (Small Subunit – 18S) nrDNA.
Additinally two loci were applied that mainly provide resolution at species
level – or even below. In addition to the abovementioned ITS regions,
also part of the β-tubulin gene was utilised, which was successfully
applied in a preliminary study on <italic>Phoma</italic> species of the section
<italic>Peyronellaea</italic> (<xref ref-type="bibr" rid="ref19">Aveskamp <italic>et
al.</italic> 2009a</xref>).</p><p>For the present study, four objectives were defined. The main objective of
this study was to reach consensus on the circumscription of the genus
<italic>Phoma</italic>. A modified definition of the genus is not only helpful in
taxonomy, but will also be of interest to plant quarantine officers
(<xref ref-type="bibr" rid="ref18">Aveskamp <italic>et al.</italic>2008</xref>). Teleomorph associations of <italic>Phoma</italic> are still
uncertain, and here we attempt to shed light on the sexual state of <italic>Phoma
s. str</italic>. Species representing all <italic>Phoma</italic> sections were included
and DNA sequences were compared with those of other species in the
<italic>Pleosporales</italic>.</p><p>Secondly, we aimed to integrate morphological and cultural features with
DNA sequence data to resolve the generic limits of taxa currently placed in
the <italic>Didymellaceae</italic>. The number of genera in this family is still
unclear. Although De Gruyter <italic>et al.</italic>(<xref ref-type="bibr" rid="ref100">2009</xref>) found a series of
genera that, according to their reconstructed phylogeny, clustered in this
family, many were not clearly defined or were morphologically distant from
each other, although all anamorph taxa found are accommodated in the
coelomycetes (<xref ref-type="bibr" rid="ref219">Sutton 1980</xref>).
Examples of these taxa were included in this study, although the number of
<italic>Ascochyta, Coniothyrium</italic> and <italic>Microsphaeropsis</italic> species is too
high to take all infrageneric taxa of these adjacent genera into account.</p><p>Further, we aimed to validate the <italic>Phoma</italic> sections, which are widely
applied in <italic>Phoma</italic> species recognition. Are the sections representing
evolutionary units, and what is the taxonomical value of the characters used
to define the sections? To judge the value of the Boeremaean taxonomic system,
representative species of all sections were studied, including the sectional
type species. The main focus was, however, to resolve the sections associated
with <italic>Didymellaceae</italic>. A single generic name, based on priority but
regardless of whether it is an “anamorph” or
“teleomorph” genus, is used for all unambiguous monophyletic
phylogenetic lineages (Crous <italic>et al</italic>.
<xref ref-type="bibr" rid="ref73">2006</xref>,
<xref ref-type="bibr" rid="ref74">2009a</xref>,
<xref ref-type="bibr" rid="ref75">b</xref>). Finally, we aimed to
assess the molecular variation within species that have historically been
placed in <italic>Phoma</italic>. Genes were tested for their potential reliability as
standard barcoding genes for <italic>Phoma</italic> species.</p><p>For this study, a sequence data set was generated and morphological data
assembled for the more than 300 well-vouchered strains available in the
culture collections of CBS (CBS-KNAW Fungal Biodiversity Centre, Utrecht, the
Netherlands) and PD (Plantenziektenkundige Dienst, Dutch Plant Protection
Service, Wageningen, the Netherlands). In addition, five species recognised in
a recent study in the section <italic>Peyronellaea</italic>(<xref ref-type="bibr" rid="ref19">Aveskamp <italic>et al.</italic>2009a</xref>) have also been included, as well as several strains that
could not be associated with any of the species that were accepted in
<italic>Phoma</italic> by Boerema <italic>et al.</italic>(<xref ref-type="bibr" rid="ref55">2004</xref>), and that were
maintained as unnamed <italic>Phoma</italic> species in the culture collections
mentioned above. These strains were recognised as taxonomic novelties and are
described at species or variety level in the present paper. Furthermore,
several species were relocated to more appropriate genera based on the results
obtained.</p></sec></sec><sec sec-type="materials|methods"><title>MATERIALS AND METHODS</title><sec><title>Strain selection</title><p>A total of 324 strains, belonging to 206 species were selected for the
present study. The majority of these species (159) belonged to the genus
<italic>Phoma</italic> or its associated teleomorphs, the remainder to genera that are
regularly confused with this genus and that belong to the
<italic>Pleosporales</italic> according to the studies published by De Gruyter <italic>et
al.</italic> (<xref ref-type="bibr" rid="ref100">2009</xref>). Besides the
anamorphous species that were included, representatives of the teleomorph
genera <italic>Didymella, Leptosphaeria, Leptosphaerulina, Macroventuria</italic> and
<italic>Pleospora</italic> were also included. The recently described genus
<italic>Atradidymella</italic> (<xref ref-type="bibr" rid="ref79">Davey & Currah
2009</xref>) was not available for study and therefore excluded.</p><p>Strains were obtained from CBS and PD culture collections in lyophilised
form or from the liquid nitrogen collection. Freeze-dried strains were revived
overnight in 2 mL malt/peptone (50 % / 50 %) liquid medium. Subsequently, the
cultures were transferred and maintained on oatmeal agar (OA,
<xref ref-type="bibr" rid="ref76">Crous <italic>et al.</italic> 2009c</xref>).
The strains that were stored at -196 °C were directly plated on the same
agar medium.</p></sec><sec><title>DNA extraction, amplification and sequence analysis</title><p>Genomic DNA extraction was performed using the Ultraclean Microbial DNA
isolation kit (Mo Bio Laboratories, Carlsbad, CA, U.S.A.), according to the
instructions of the manufacturer. All DNA extracts were diluted 10 × in
milliQ water and stored at 4 °C before their use as PCR templates.</p><p>For nucleotide sequence comparisons fragments of four loci were analysed:
LSU, SSU, ITS, and TUB. Amplification of LSU and SSU was conducted utilising
the primer combination LR0R (<xref ref-type="bibr" rid="ref180">Rehner &
Samuels 1994</xref>) and LR7
(<xref ref-type="bibr" rid="ref229">Vilgalys & Hester 1990</xref>)
for LSU sequencing and the primer pair NS1 and NS4
(<xref ref-type="bibr" rid="ref234">White <italic>et al.</italic> 1990</xref>)
for SSU. The PCRs were performed in a 2720 Thermal Cycler (Applied Biosystems,
Foster City, California) in a total volume of 12.5 μL. The PCR mixture
contained 0.5 μL 10 × diluted genomic DNA, 0.2 μM of each primer,
0.5 Unit Taq polymerase E (Genaxxon Bioscience, Germany), 0.04 mM (SSU) or
0.06 mM (LSU) of each of the dNTP, 2 mM MgCl<sub>2</sub> and 1 × PCR
buffer E incomplete (Genaxxon Bioscience). Conditions for amplification for
both regions were an initial denaturation step of 5 min at 94 °C, followed
by 35 cycles of denaturation, annealing and elongation and a final elongation
step of 7 min at 72 °C. For the SSU amplification, the 35 cycles consisted
of 30 s at 94 °C, 50 s at 48 °C and 90 s at 72 °C; for the LSU 45
s at 94 °C, 45 s at 48 °C and 2 min at 72 °C. The loci ITS and TUB
were amplified as described by Aveskamp <italic>et al</italic>.
(<xref ref-type="bibr" rid="ref19">2009a</xref>), using the primer
pairs V9G (<xref ref-type="bibr" rid="ref107">De Hoog & Gerrits van den
Ende 1998</xref>) and ITS4 (<xref ref-type="bibr" rid="ref234">White
<italic>et al.</italic> 1990</xref>) for ITS sequencing and the BT2Fw and BT4Rd
primer pair (<xref ref-type="bibr" rid="ref19">Aveskamp <italic>et al.</italic>2009a</xref>) for sequencing of the TUB locus. PCR products were analysed
by electrophoresis in a 1.0 % (w/v) agarose gel containing 0.1 ug/mL ethidium
bromide in 1 × TAE buffer (0.4 M Tris, 0.05 M glacial acetetic acid 0.01
M ethylenediamine tetraacetic acid [EDTA], pH 7.85). The amplicons were
visualised under UV light. Hyperladder I (Bioline, Luckenwalde, Germany) was
applied as size standard.</p><p>The obtained amplicons were sequenced in both directions using the same
primer combinations, except for LSU, where an additional primer, LR5
(<xref ref-type="bibr" rid="ref234">White <italic>et al.</italic> 1990</xref>)
was further required to assure complete coverage of the locus. Sequencing
reactions were prepared with the BigDye terminator chemistry v. 3.1 (Applied
Biosystems) according to the manufacturer's recommendations. Sequence products
were purified with Sephadex G-50 Fine (Amersham Biosciences, Roosendaal, the
Netherlands) and subsequently separated and analysed on an ABI Prism 3730 DNA
Sequencer (Applied Biosystems). Consensus sequences were computed from the
forward and reverse sequences using the BioNumerics v. 4.61 software package
(Applied Maths, St-Martens-Latem, Belgium). The consensus sequences are
deposited in GenBank (For GenBank accession numbers see Tables
<xref ref-type="table" rid="tbl2">2</xref>,
<xref ref-type="table" rid="tbl3">3</xref>).</p><p><table-wrap position="float" id="tbl2"><label>Table 2.</label><caption><p>Isolates of <italic>Phoma</italic> and related genera used for DNA analyses. The
GenBank accession numbers in bold have been obtained from other studies.</p></caption><table frame="hsides" rules="groups"><thead><tr><th align="left" valign="top"><bold>Strain
no.</bold><xref ref-type="table-fn" rid="tblfn2"><bold><sup>1</sup></bold></xref></th><th align="center" valign="top"><bold>Holomorph</bold><xref ref-type="table-fn" rid="tblfn3"><bold><sup>2</sup></bold></xref></th><th valign="top" colspan="2" align="center"><bold>GenBank no.</bold><hr></hr></th><th align="center" valign="top"><bold>Original substrate</bold></th><th align="center" valign="top"><bold>Locality</bold></th></tr><tr><th valign="top" align="left"></th><th valign="top" align="center"></th><th valign="top" align="center"><bold>SSU</bold></th><th valign="top" align="center"><bold>LSU</bold></th><th valign="top" align="center"></th><th valign="top" align="center"></th></tr></thead><tbody><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=129.79&link_type=cbs">CBS 129.79</ext-link></td><td align="left" valign="top"><italic>Ampelomyces quisqualis</italic></td><td align="center" valign="top"><bold>EU754029</bold></td><td align="center" valign="top"><bold>EU754128</bold></td><td align="left" valign="top"> Mildew on <italic>Cucumis sativus</italic></td><td align="left" valign="top"> Canada
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=543.70&link_type=cbs">CBS 543.70</ext-link></td><td align="left" valign="top"><italic>Aposphaeria populina</italic></td><td align="center" valign="top"><bold>EU754031</bold></td><td align="center" valign="top"><bold>EU754130</bold></td><td align="left" valign="top"><italic>Populus canadensis</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=246.79&link_type=cbs">CBS 246.79</ext-link>; PD
77/655
</td><td align="left" valign="top"><italic>Ascochyta caulina</italic> T
</td><td align="center" valign="top"><bold>EU754032</bold></td><td align="center" valign="top"><bold>EU754131</bold></td><td align="left" valign="top"><italic>Atriplex hastata</italic></td><td align="left" valign="top"> Germany
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=544.74&link_type=cbs">CBS 544.74</ext-link></td><td align="left" valign="top"><italic>Ascochyta hordei</italic> var. <italic>hordei</italic></td><td align="center" valign="top"><bold>EU754035</bold></td><td align="center" valign="top"><bold>EU754134</bold></td><td align="left" valign="top"><italic>Triticum aevestum</italic></td><td align="left" valign="top"> South Africa
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=117477&link_type=cbs">CBS 117477</ext-link></td><td align="left" valign="top"><italic>Ascochyta</italic> sp.
</td><td align="center" valign="top"> GU238202
</td><td align="center" valign="top"> GU237926
</td><td align="left" valign="top"><italic>Salicornia australis</italic></td><td align="left" valign="top"> New Zealand
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=265.94&link_type=cbs">CBS 265.94</ext-link></td><td align="left" valign="top"><italic>Asteromella tiliae</italic></td><td align="center" valign="top"><bold>EU754040</bold></td><td align="center" valign="top"><bold>EU754139</bold></td><td align="left" valign="top"><italic>Tilia platyphilos</italic></td><td align="left" valign="top"> Austria
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=431.74&link_type=cbs">CBS 431.74</ext-link>; PD
74/2447
</td><td align="left" valign="top"><italic>Boeremia exigua</italic> var. <italic>exigua</italic> B
</td><td align="center" valign="top"><bold>EU754084</bold></td><td align="center" valign="top"><bold>EU754183</bold></td><td align="left" valign="top"><italic>Solanum tuberosum</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=341.67&link_type=cbs">CBS 341.67</ext-link>; CECT
20055; IMI 331912
</td><td align="left" valign="top"><italic>Boeremia foveata</italic> B
</td><td align="center" valign="top"> GU238203
</td><td align="center" valign="top"> GU237947
</td><td align="left" valign="top"><italic>Solanum tuberosum</italic></td><td align="left" valign="top"> U.K.
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=148.94&link_type=cbs">CBS 148.94</ext-link></td><td align="left" valign="top"><italic>Chaetasbolisia erysiphoides</italic></td><td align="center" valign="top"><bold>EU754041</bold></td><td align="center" valign="top"><bold>EU754140</bold></td><td align="left" valign="top"> Unknown
</td><td align="left" valign="top"> Unknown
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=216.75&link_type=cbs">CBS 216.75</ext-link>; PD
71/1030
</td><td align="left" valign="top"><italic>Chaetosphaeronema hispidulum</italic></td><td align="center" valign="top"><bold>EU754045</bold></td><td align="center" valign="top"><bold>EU754144</bold></td><td align="left" valign="top"><italic>Anthyllis vulneraria</italic></td><td align="left" valign="top"> Germany
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=589.79&link_type=cbs">CBS 589.79</ext-link></td><td align="left" valign="top"><italic>Coniothyrium concentricum</italic></td><td align="center" valign="top"><bold>EU754053</bold></td><td align="center" valign="top"><bold>EU754152</bold></td><td align="left" valign="top"><italic>Yucca</italic> sp.
</td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=797.95&link_type=cbs">CBS 797.95</ext-link></td><td align="left" valign="top"><italic>Coniothyrium fuckelii</italic></td><td align="center" valign="top"> GU238204
</td><td align="center" valign="top"> GU237960
</td><td align="left" valign="top"><italic>Rubus</italic> sp.
</td><td align="left" valign="top"> Denmark
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=400.71&link_type=cbs">CBS 400.71</ext-link></td><td align="left" valign="top"><italic>Coniothyrium palmarum</italic></td><td align="center" valign="top"><bold>EU754054</bold></td><td align="center" valign="top"><bold>EU754153</bold></td><td align="left" valign="top"><italic>Chamaerops humilis</italic></td><td align="left" valign="top"> Italy
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=122787&link_type=cbs">CBS 122787</ext-link>; PD
03486691
</td><td align="left" valign="top"><italic>Coniothyrium</italic> sp.
</td><td align="center" valign="top"><bold>EU754052</bold></td><td align="center" valign="top"><bold>EU754151</bold></td><td align="left" valign="top"> Unknown
</td><td align="left" valign="top"> Germany
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=183.55&link_type=cbs">CBS 183.55</ext-link></td><td align="left" valign="top"><italic>Didymella exigua</italic> T
</td><td align="center" valign="top"><bold>EU754056</bold></td><td align="center" valign="top"><bold>EU754155</bold></td><td align="left" valign="top"><italic>Rumex arifolius</italic></td><td align="left" valign="top"> France
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=524.77&link_type=cbs">CBS 524.77</ext-link></td><td align="left" valign="top"><italic>Didymella fabae</italic></td><td align="center" valign="top"><bold>EU754034</bold></td><td align="center" valign="top"><bold>EU754133</bold></td><td align="left" valign="top"><italic>Phaseolus vulgaris</italic></td><td align="left" valign="top"> Belgium
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=581.83&link_type=cbs">CBS 581.83</ext-link>A
</td><td align="left" valign="top"><italic>Didymella rabiei</italic></td><td align="center" valign="top"> GU238205
</td><td align="center" valign="top"> GU237970
</td><td align="left" valign="top"><italic>Cicer arietinum</italic></td><td align="left" valign="top"> Syria
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=173.73&link_type=cbs">CBS 173.73</ext-link>; ATCC
24428; IMI 164070
</td><td align="left" valign="top"><italic>Epicoccum nigrum</italic> T
</td><td align="center" valign="top"> GU238206
</td><td align="center" valign="top"> GU237975
</td><td align="left" valign="top"><italic>Dactylis glomerata</italic></td><td align="left" valign="top"> U.S.A.
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=298.36&link_type=cbs">CBS 298.36</ext-link></td><td align="left" valign="top"><italic>Leptosphaeria biglobosa</italic></td><td align="center" valign="top"> GU238207
</td><td align="center" valign="top"> GU237980
</td><td align="left" valign="top"><italic>Brassica napus</italic> var. <italic>napobrassica</italic></td><td align="left" valign="top"> Unknown
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=127.23&link_type=cbs">CBS 127.23</ext-link>; MUCL
9930
</td><td align="left" valign="top"><italic>Leptosphaeria maculans</italic></td><td align="center" valign="top"><bold>EU754090</bold></td><td align="center" valign="top"><bold>EU754189</bold></td><td align="left" valign="top"><italic>Brassica</italic> sp.
</td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=939.69&link_type=cbs">CBS 939.69</ext-link></td><td align="left" valign="top"><italic>Leptosphaerulina australis</italic></td><td align="center" valign="top"><bold>EU754068</bold></td><td align="center" valign="top"><bold>EU754167</bold></td><td align="left" valign="top"> Soil
</td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=525.71&link_type=cbs">CBS 525.71</ext-link></td><td align="left" valign="top"><italic>Macroventuria anomochaeta</italic> T
</td><td align="center" valign="top"> GU238208
</td><td align="center" valign="top"> GU237984
</td><td align="left" valign="top"> Decayed canvas
</td><td align="left" valign="top"> South Africa
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=442.83&link_type=cbs">CBS 442.83</ext-link></td><td align="left" valign="top"><italic>Microsphaeropsis olivacea</italic></td><td align="center" valign="top"><bold>EU754072</bold></td><td align="center" valign="top"><bold>EU754171</bold></td><td align="left" valign="top"><italic>Taxus baccata</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=331.37&link_type=cbs">CBS 331.37</ext-link></td><td align="left" valign="top"><italic>Neottiosporina paspali</italic></td><td align="center" valign="top"><bold>EU754073</bold></td><td align="center" valign="top"><bold>EU754172</bold></td><td align="left" valign="top"><italic>Paspalum notatum</italic></td><td align="left" valign="top"> U.S.A.
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=122786&link_type=cbs">CBS 122786</ext-link>; PD
99/1064-1
</td><td align="left" valign="top"><italic>Paraconiothyrium minitans</italic></td><td align="center" valign="top"><bold>EU754075</bold></td><td align="center" valign="top"><bold>EU754174</bold></td><td align="left" valign="top"> Unknown
</td><td align="left" valign="top"> Unknown
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=626.68&link_type=cbs">CBS 626.68</ext-link>; IMI
108771
</td><td align="left" valign="top"><italic>Peyronellaea gardeniae</italic> T
</td><td align="center" valign="top"><bold>GQ387534</bold></td><td align="center" valign="top"><bold>GQ387595</bold></td><td align="left" valign="top"><italic>Gardenia jasminoides</italic></td><td align="left" valign="top"> India
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=528.66&link_type=cbs">CBS 528.66</ext-link>; PD
63/590
</td><td align="left" valign="top"><italic>Peyronellaea glomerata</italic> B
</td><td align="center" valign="top"><bold>EU754085</bold></td><td align="center" valign="top"><bold>EU754184</bold></td><td align="left" valign="top"><italic>Chrysanthemum</italic> sp.
</td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=531.66&link_type=cbs">CBS 531.66</ext-link></td><td align="left" valign="top"><italic>Peyronellaea pinodella</italic> B
</td><td align="center" valign="top"> GU238209
</td><td align="center" valign="top"> GU238017
</td><td align="left" valign="top"><italic>Trifolium pratense</italic></td><td align="left" valign="top"> U.S.A.
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=235.55&link_type=cbs">CBS 235.55</ext-link></td><td align="left" valign="top"><italic>Peyronellaea pinodes</italic></td><td align="center" valign="top"> GU238210
</td><td align="center" valign="top"> GU238021
</td><td align="left" valign="top"> Unknown
</td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=588.69&link_type=cbs">CBS 588.69</ext-link></td><td align="left" valign="top"><italic>Peyronellaea zeae-maydis</italic> T
</td><td align="center" valign="top"><bold>EU754093</bold></td><td align="center" valign="top"><bold>EU754192</bold></td><td align="left" valign="top"><italic>Zea mays</italic></td><td align="left" valign="top"> U.S.A.
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=110110&link_type=cbs">CBS 110110</ext-link></td><td align="left" valign="top"><italic>Phaeosphaeria oryzae</italic></td><td align="center" valign="top"><bold>GQ387530</bold></td><td align="center" valign="top"><bold>GQ387591</bold></td><td align="left" valign="top"><italic>Oryza sativa</italic></td><td align="left" valign="top"> South Korea
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=297.74&link_type=cbs">CBS 297.74</ext-link></td><td align="left" valign="top"><italic>Phialophorophoma litoralis</italic></td><td align="center" valign="top"><bold>EU754078</bold></td><td align="center" valign="top"><bold>EU754177</bold></td><td align="left" valign="top"> Sea water
</td><td align="left" valign="top"> Montenegro
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=285.72&link_type=cbs">CBS 285.72</ext-link></td><td align="left" valign="top"><italic>Phoma apiicola</italic> B
</td><td align="center" valign="top"> GU238211
</td><td align="center" valign="top"> GU238040
</td><td align="left" valign="top"><italic>Apium graveolens</italic> var. <italic>rapaceum</italic></td><td align="left" valign="top"> Germany
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=337.65&link_type=cbs">CBS 337.65</ext-link>; ATCC
16195; IMI 113693
</td><td align="left" valign="top"><italic>Phoma capitulum</italic> B
</td><td align="center" valign="top"> GU238212
</td><td align="center" valign="top"> GU238054
</td><td align="left" valign="top"> Soil
</td><td align="left" valign="top"> India
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=522.66&link_type=cbs">CBS 522.66</ext-link></td><td align="left" valign="top"><italic>Phoma chrysanthemicola</italic> T
</td><td align="center" valign="top"><bold>GQ387521</bold></td><td align="center" valign="top"><bold>GQ387582</bold></td><td align="left" valign="top"><italic>Chrysanthemum morifolium</italic></td><td align="left" valign="top"> U.K.
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=100311&link_type=cbs">CBS 100311</ext-link></td><td align="left" valign="top"><italic>Phoma complanata</italic></td><td align="center" valign="top"><bold>EU754082</bold></td><td align="center" valign="top"><bold>EU754181</bold></td><td align="left" valign="top"><italic>Heracleum sphondylium</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=345.78&link_type=cbs">CBS 345.78</ext-link>; PD
76/1015
</td><td align="left" valign="top"><italic>Phoma dimorphospora</italic></td><td align="center" valign="top"> GU238213
</td><td align="center" valign="top"> GU238069
</td><td align="left" valign="top"><italic>Chenopodium quinoa</italic></td><td align="left" valign="top"> Peru
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=527.66&link_type=cbs">CBS 527.66</ext-link></td><td align="left" valign="top"><italic>Phoma eupyrena</italic> B
</td><td align="center" valign="top"> GU238214
</td><td align="center" valign="top"> GU238072
</td><td align="left" valign="top"> Soil
</td><td align="left" valign="top"> Germany
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=161.78&link_type=cbs">CBS 161.78</ext-link></td><td align="left" valign="top"><italic>Phoma fallens</italic> B
</td><td align="center" valign="top"> GU238215
</td><td align="center" valign="top"> GU238074
</td><td align="left" valign="top"><italic>Olea europaea</italic></td><td align="left" valign="top"> New Zealand
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=170.70&link_type=cbs">CBS 170.70</ext-link>; ATCC
22707; CECT 20011; IMI 163514; PD 70/Alk
</td><td align="left" valign="top"><italic>Phoma fimeti</italic> T
</td><td align="center" valign="top"><bold>GQ387523</bold></td><td align="center" valign="top"><bold>GQ387584</bold></td><td align="left" valign="top"><italic>Apium graveolens</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=178.93&link_type=cbs">CBS 178.93</ext-link>; PD
82/1062
</td><td align="left" valign="top"><italic>Phoma flavescens</italic> T
</td><td align="center" valign="top"> GU238216
</td><td align="center" valign="top"> GU238075
</td><td align="left" valign="top"> Soil
</td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=314.80&link_type=cbs">CBS 314.80</ext-link></td><td align="left" valign="top"><italic>Phoma flavigena</italic> T
</td><td align="center" valign="top"> GU238217
</td><td align="center" valign="top"> GU238076
</td><td align="left" valign="top"> Water
</td><td align="left" valign="top"> Romania
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=633.92&link_type=cbs">CBS 633.92</ext-link>; ATCC
36786; VKM MF-325
</td><td align="left" valign="top"><italic>Phoma fungicola</italic></td><td align="center" valign="top"><bold>EU754028</bold></td><td align="center" valign="top"><bold>EU754127</bold></td><td align="left" valign="top"><italic>Microsphaera alphitoides</italic> on <italic>Quercus</italic> sp.
</td><td align="left" valign="top"> Ukraine
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=284.70&link_type=cbs">CBS 284.70</ext-link></td><td align="left" valign="top"><italic>Phoma glaucispora</italic> B
</td><td align="center" valign="top"> GU238218
</td><td align="center" valign="top"> GU238078
</td><td align="left" valign="top"><italic>Nerium oleander</italic></td><td align="left" valign="top"> Italy
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=175.93&link_type=cbs">CBS 175.93</ext-link>; PD
92/370
</td><td align="left" valign="top"><italic>Phoma haematocycla</italic> T
</td><td align="center" valign="top"> GU238219
</td><td align="center" valign="top"> GU238080
</td><td align="left" valign="top"><italic>Phormium tenax</italic></td><td align="left" valign="top"> New Zealand
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=615.75&link_type=cbs">CBS 615.75</ext-link>; PD
73/665, IMI 199779
</td><td align="left" valign="top"><italic>Phoma herbarum</italic> B
</td><td align="center" valign="top"><bold>EU754087</bold></td><td align="center" valign="top"><bold>EU754186</bold></td><td align="left" valign="top"><italic>Rosa multiflora</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=448.68&link_type=cbs">CBS 448.68</ext-link></td><td align="left" valign="top"><italic>Phoma heteromorphospora</italic> B
</td><td align="center" valign="top"><bold>EU754088</bold></td><td align="center" valign="top"><bold>EU754187</bold></td><td align="left" valign="top"><italic>Chenopodium album</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=467.76&link_type=cbs">CBS 467.76</ext-link></td><td align="left" valign="top"><italic>Phoma incompta</italic> B
</td><td align="center" valign="top"> GU238220
</td><td align="center" valign="top"> GU238087
</td><td align="left" valign="top"><italic>Olea europaea</italic></td><td align="left" valign="top"> Greece
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=253.92&link_type=cbs">CBS 253.92</ext-link>; PD
70/998
</td><td align="left" valign="top"><italic>Phoma lini</italic> B
</td><td align="center" valign="top"> GU238221
</td><td align="center" valign="top"> GU238093
</td><td align="left" valign="top"> Water
</td><td align="left" valign="top"> U.S.A.
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=529.66&link_type=cbs">CBS 529.66</ext-link>; PD
66/521
</td><td align="left" valign="top"><italic>Phoma macrostoma</italic> var. <italic>macrostoma</italic> B
</td><td align="center" valign="top"> GU238222
</td><td align="center" valign="top"> GU238098
</td><td align="left" valign="top"><italic>Malus sylvestris</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=316.90&link_type=cbs">CBS 316.90</ext-link></td><td align="left" valign="top"><italic>Phoma medicaginis</italic> var. <italic>medicaginis</italic></td><td align="center" valign="top"> GU238223
</td><td align="center" valign="top"> GU238103
</td><td align="left" valign="top"><italic>Medicago sativa</italic></td><td align="left" valign="top"> Czech Republic
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=509.91&link_type=cbs">CBS 509.91</ext-link>; PD
77/920
</td><td align="left" valign="top"><italic>Phoma minutispora</italic></td><td align="center" valign="top"> GU238224
</td><td align="center" valign="top"> GU238108
</td><td align="left" valign="top"> Saline soil
</td><td align="left" valign="top"> India
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=501.91&link_type=cbs">CBS 501.91</ext-link>; PD
83/888
</td><td align="left" valign="top"><italic>Phoma multipora</italic> B
</td><td align="center" valign="top"> GU238225
</td><td align="center" valign="top"> GU238109
</td><td align="left" valign="top"> Unknown
</td><td align="left" valign="top"> Egypt
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=376.91&link_type=cbs">CBS 376.91</ext-link>;
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=328.78&link_type=cbs">CBS 328.78</ext-link>, PD
77/1177
</td><td align="left" valign="top"><italic>Phoma opuntiae</italic> B
</td><td align="center" valign="top"> GU238226
</td><td align="center" valign="top"> GU238123
</td><td align="left" valign="top"><italic>Opuntia ficus-indica</italic>.
</td><td align="left" valign="top"> Peru
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=560.81&link_type=cbs">CBS 560.81</ext-link>; PD
92/1569; PDDCC 6614
</td><td align="left" valign="top"><italic>Phoma paspali</italic> T
</td><td align="center" valign="top"> GU238227
</td><td align="center" valign="top"> GU238124
</td><td align="left" valign="top"><italic>Paspalum dilatatum</italic></td><td align="left" valign="top"> New Zealand
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=445.81&link_type=cbs">CBS 445.81</ext-link>; PDDCC
7049
</td><td align="left" valign="top"><italic>Phoma pratorum</italic> T
</td><td align="center" valign="top"> GU238228
</td><td align="center" valign="top"> GU238136
</td><td align="left" valign="top"><italic>Lolium perenne</italic></td><td align="left" valign="top"> New Zealand
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=111.79&link_type=cbs">CBS 111.79</ext-link>; PD
76/437; IMI 386094
</td><td align="left" valign="top"><italic>Phoma radicina</italic> B
</td><td align="center" valign="top"><bold>EU754092</bold></td><td align="center" valign="top"><bold>EU754191</bold></td><td align="left" valign="top"><italic>Malus sylvestris</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=138.96&link_type=cbs">CBS 138.96</ext-link>; PD
82/653
</td><td align="left" valign="top"><italic>Phoma samarorum</italic> B
</td><td align="center" valign="top"><bold>GQ387517</bold></td><td align="center" valign="top"><bold>GQ387578</bold></td><td align="left" valign="top"><italic>Phlox paniculata</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=343.85&link_type=cbs">CBS 343.85</ext-link>; IMI
386097
</td><td align="left" valign="top"><italic>Phoma terricola</italic> T
</td><td align="center" valign="top"><bold>GQ387563</bold></td><td align="center" valign="top"><bold>GQ387624</bold></td><td align="left" valign="top"><italic>Globodera pallida</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=630.68&link_type=cbs">CBS 630.68</ext-link>; PD
68/141
</td><td align="left" valign="top"><italic>Phoma valerianae</italic> B
</td><td align="center" valign="top"> GU238229
</td><td align="center" valign="top"> GU238150
</td><td align="left" valign="top"><italic>Valeriana phu</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=539.63&link_type=cbs">CBS 539.63</ext-link></td><td align="left" valign="top"><italic>Phoma vasinfecta</italic> T
</td><td align="center" valign="top"> GU238230
</td><td align="center" valign="top"> GU238151
</td><td align="left" valign="top"><italic>Chrysanthemum</italic> sp.
</td><td align="left" valign="top"> Greece
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=306.68&link_type=cbs">CBS 306.68</ext-link></td><td align="left" valign="top"><italic>Phoma violicola</italic> B
</td><td align="center" valign="top"> GU238231
</td><td align="center" valign="top"> GU238156
</td><td align="left" valign="top"><italic>Viola tricolor</italic></td><td align="left" valign="top"> Unknown
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=523.66&link_type=cbs">CBS 523.66</ext-link>; PD
66/270
</td><td align="left" valign="top"><italic>Pleospora betae</italic> B
</td><td align="center" valign="top"><bold>EU754080</bold></td><td align="center" valign="top"><bold>EU754179</bold></td><td align="left" valign="top"><italic>Beta vulgaris</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=191.86&link_type=cbs">CBS 191.86</ext-link>; IMI
276975
</td><td align="left" valign="top"><italic>Pleospora herbarum</italic> T
</td><td align="center" valign="top"> GU238232
</td><td align="center" valign="top"> GU238160
</td><td align="left" valign="top"><italic>Medicago sativa</italic></td><td align="left" valign="top"> India
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=257.68&link_type=cbs">CBS 257.68</ext-link>; IMI
331911
</td><td align="left" valign="top"><italic>Pleurophoma cava</italic></td><td align="center" valign="top"><bold>EU754100</bold></td><td align="center" valign="top"><bold>EU754199</bold></td><td align="left" valign="top"> Soil
</td><td align="left" valign="top"> Germany
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=398.61&link_type=cbs">CBS 398.61</ext-link>; IMI
070678
</td><td align="left" valign="top"><italic>Pseudorobillarda phragmitis</italic> T
</td><td align="center" valign="top"><bold>EU754104</bold></td><td align="center" valign="top"><bold>EU754203</bold></td><td align="left" valign="top"><italic>Phragmites australis</italic></td><td align="left" valign="top"> U.K.
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=122789&link_type=cbs">CBS 122789</ext-link>; PD
03486800
</td><td align="left" valign="top"><italic>Pyrenochaeta acicola</italic></td><td align="center" valign="top"><bold>EU754105</bold></td><td align="center" valign="top"><bold>EU754204</bold></td><td align="left" valign="top"><italic>Hordeum vulgare</italic></td><td align="left" valign="top"> Unknown
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=306.65&link_type=cbs">CBS 306.65</ext-link></td><td align="left" valign="top"><italic>Pyrenochaeta lycopersici</italic> T
</td><td align="center" valign="top"><bold>EU754106</bold></td><td align="center" valign="top"><bold>EU754205</bold></td><td align="left" valign="top"><italic>Lycopersicon esculentum</italic></td><td align="left" valign="top"> Germany
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=407.76&link_type=cbs">CBS 407.76</ext-link></td><td align="left" valign="top"><italic>Pyrenochaeta nobilis</italic> T
</td><td align="center" valign="top"><bold>EU754107</bold></td><td align="center" valign="top"><bold>EU754206</bold></td><td align="left" valign="top"><italic>Laurus nobilis</italic></td><td align="left" valign="top"> Italy
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=252.60&link_type=cbs">CBS 252.60</ext-link>; ATCC
13735
</td><td align="left" valign="top"><italic>Pyrenochaeta romeroi</italic> T
</td><td align="center" valign="top"><bold>EU754108</bold></td><td align="center" valign="top"><bold>EU754209</bold></td><td align="left" valign="top"> Man
</td><td align="left" valign="top"> Venezuela
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=524.50&link_type=cbs">CBS 524.50</ext-link></td><td align="left" valign="top"><italic>Sporormiella minima</italic></td><td align="center" valign="top"><bold>DQ678003</bold></td><td align="center" valign="top"><bold>DQ678056</bold></td><td align="left" valign="top"> Goat dung
</td><td align="left" valign="top"> Panama
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=343.86&link_type=cbs">CBS 343.86</ext-link></td><td align="left" valign="top"><italic>Stagonospora neglecta</italic> var. <italic>colorata</italic></td><td align="center" valign="top"><bold>EU754119</bold></td><td align="center" valign="top"><bold>EU754218</bold></td><td align="left" valign="top"><italic>Phragmites australis</italic></td><td align="left" valign="top"> France
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=101.80&link_type=cbs">CBS 101.80</ext-link>; PD
75/909; IMI 386090
</td><td align="left" valign="top"><italic>Stagonosporopsis andigena</italic> B
</td><td align="center" valign="top"> GU238233
</td><td align="center" valign="top"> GU238169
</td><td align="left" valign="top"><italic>Solanum</italic> sp.
</td><td align="left" valign="top"> Peru
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=133.96&link_type=cbs">CBS 133.96</ext-link>; PD
79/127
</td><td align="left" valign="top"><italic>Stagonosporopsis cucurbitacearum</italic></td><td align="center" valign="top"> GU238234
</td><td align="center" valign="top"> GU238181
</td><td align="left" valign="top"><italic>Cucurbita</italic> sp.
</td><td align="left" valign="top"> New Zealand
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=631.68&link_type=cbs">CBS 631.68</ext-link>; PD
68/147
</td><td align="left" valign="top"><italic>Stagonosporopsis dennisii</italic> B
</td><td align="center" valign="top"> GU238235
</td><td align="center" valign="top"> GU238182
</td><td align="left" valign="top"><italic>Solidago floribunda</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=164.31&link_type=cbs">CBS 164.31</ext-link></td><td align="left" valign="top"><italic>Stenocarpella macrospora</italic></td><td align="center" valign="top"><bold>EU754121</bold></td><td align="center" valign="top"><bold>EU754220</bold></td><td align="left" valign="top"><italic>Zea mays</italic></td><td align="left" valign="top">Unknown
</td></tr></tbody></table><table-wrap-foot><fn id="tblfn2"><label>1</label><p>ATCC: American Type Culture Collection, Virginia, U.S.A.; CBS:
Centraalbureau voor Schimmelcultures, Utrecht, The Netherlands; CECT:
Colección Española de Cultivos Tipo, Valencia University, Spain;
IMI: International Mycological Institute, CABI-Bioscience, Egham, Bakeham
Lane, U.K.; MUCL: Mycotheque de l'Universite catholique de Louvain,
Louvain-la-Neuve, Belgium; PD: Plant Protection Service, Wageningen, the
Netherlands; PDDCC: Plant Diseases Division Culture Collection, Auckland, New
Zealand; VKM: All-Russian Collection of Microorganisms, Pushchino, Russia.</p></fn><fn id="tblfn3"><label>2</label><p>T: Ex-type strain; B: Reference strain according to Boerema <italic>et al.</italic>(<xref ref-type="bibr" rid="ref55">2004</xref>).</p></fn></table-wrap-foot></table-wrap></p><p><table-wrap position="float" id="tbl3"><label>Table 3.</label><caption><p>Strains from the <italic>Didymellaceae</italic> used for DNA analyses. The GenBank
accession numbers in bold have been obtained from other studies.</p></caption><table frame="hsides" rules="groups"><thead><tr><th align="left" valign="top"><bold>Strain
no.</bold><xref ref-type="table-fn" rid="tblfn4"><bold>1</bold></xref></th><th align="center" valign="top"><bold>Holomorph</bold><xref ref-type="table-fn" rid="tblfn5"><bold>2</bold></xref></th><th valign="top" colspan="3" align="center"><bold>GenBank no.</bold><hr></hr></th><th align="center" valign="top"><bold>Original substrate</bold></th><th align="center" valign="top"><bold>Locality</bold></th></tr><tr><th valign="top" align="left"></th><th valign="top" align="center"></th><th valign="top" align="center"><bold>LSU</bold></th><th valign="top" align="center"><bold>ITS</bold></th><th valign="top" align="center"><bold>TUB</bold></th><th valign="top" align="center"></th><th valign="top" align="center"></th></tr></thead><tbody><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=544.74&link_type=cbs">CBS 544.74</ext-link></td><td align="left" valign="top"><italic>Ascochyta hordei</italic> var. <italic>hordei</italic></td><td align="center" valign="top"><bold>EU754134</bold></td><td align="center" valign="top"> GU237887
</td><td align="center" valign="top"> GU237488
</td><td align="left" valign="top"><italic>Triticum aevestum</italic></td><td align="left" valign="top"> South Africa
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=109.79&link_type=cbs">CBS 109.79</ext-link>; PD
77/747
</td><td align="left" valign="top"><italic>Boeremia crinicola</italic> B
</td><td align="center" valign="top"> GU237927
</td><td align="center" valign="top"> GU237737
</td><td align="center" valign="top"> GU237489
</td><td align="left" valign="top"><italic>Crinum powellii</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=118.93&link_type=cbs">CBS 118.93</ext-link>; PD
70/195
</td><td align="left" valign="top"><italic>Boeremia crinicola</italic></td><td align="center" valign="top"> GU237928
</td><td align="center" valign="top"> GU237758
</td><td align="center" valign="top"> GU237490
</td><td align="left" valign="top"><italic>Crinum</italic> sp.
</td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=101194&link_type=cbs">CBS 101194</ext-link>; PD
79/687; IMI 373349
</td><td align="left" valign="top"><italic>Boeremia diversispora</italic></td><td align="center" valign="top"> GU237929
</td><td align="center" valign="top"> GU237716
</td><td align="center" valign="top"> GU237491
</td><td align="left" valign="top"><italic>Phaseolus vulgaris</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=102.80&link_type=cbs">CBS 102.80</ext-link>; PD
79/61; CECT 20049; IMI 331907
</td><td align="left" valign="top"><italic>Boeremia diversispora</italic> B
</td><td align="center" valign="top"> GU237930
</td><td align="center" valign="top"> GU237725
</td><td align="center" valign="top"> GU237492
</td><td align="left" valign="top"><italic>Phaseolus vulgaris</italic></td><td align="left" valign="top"> Kenya
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=119730&link_type=cbs">CBS 119730</ext-link></td><td align="left" valign="top"><italic>Boeremia exigua</italic> var. <italic>coffeae</italic></td><td align="center" valign="top"> GU237942
</td><td align="center" valign="top"> GU237759
</td><td align="center" valign="top"> GU237504
</td><td align="left" valign="top"><italic>Coffea arabica</italic></td><td align="left" valign="top"> Brazil
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=109183&link_type=cbs">CBS 109183</ext-link>; IMI
300060; PD 2000/10506
</td><td align="left" valign="top"><italic>Boeremia exigua</italic> var. <italic>coffeae</italic> B
</td><td align="center" valign="top"> GU237943
</td><td align="center" valign="top"> GU237748
</td><td align="center" valign="top"> GU237505
</td><td align="left" valign="top"><italic>Coffea arabica</italic></td><td align="left" valign="top"> Cameroon
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=431.74&link_type=cbs">CBS 431.74</ext-link>; PD
74/2447
</td><td align="left" valign="top"><italic>Boeremia exigua</italic> var. <italic>exigua</italic> B
</td><td align="center" valign="top"><bold>EU754183</bold></td><td align="center" valign="top"><bold>FJ427001</bold></td><td align="center" valign="top"><bold>FJ427112</bold></td><td align="left" valign="top"><italic>Solanum tuberosum</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=101150&link_type=cbs">CBS 101150</ext-link>; PD
79/118
</td><td align="left" valign="top"><italic>Boeremia exigua</italic> var. <italic>exigua</italic></td><td align="center" valign="top"> GU237933
</td><td align="center" valign="top"> GU237715
</td><td align="center" valign="top"> GU237495
</td><td align="left" valign="top"><italic>Cichorium intybus</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=101197&link_type=cbs">CBS 101197</ext-link>; PD
95/721
</td><td align="left" valign="top"><italic>Boeremia exigua</italic> var. <italic>forsythiae</italic></td><td align="center" valign="top"> GU237931
</td><td align="center" valign="top"> GU237718
</td><td align="center" valign="top"> GU237493
</td><td align="left" valign="top"><italic>Forsythia</italic> sp.
</td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=101213&link_type=cbs">CBS 101213</ext-link>; PD
92/959
</td><td align="left" valign="top"><italic>Boeremia exigua</italic> var. <italic>forsythiae</italic> B
</td><td align="center" valign="top"> GU237932
</td><td align="center" valign="top"> GU237723
</td><td align="center" valign="top"> GU237494
</td><td align="left" valign="top"><italic>Forsythia</italic> sp.
</td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=101196&link_type=cbs">CBS 101196</ext-link>; PD
79/176
</td><td align="left" valign="top"><italic>Boeremia exigua</italic> var. <italic>heteromorpha</italic></td><td align="center" valign="top"> GU237934
</td><td align="center" valign="top"> GU237717
</td><td align="center" valign="top"> GU237496
</td><td align="left" valign="top"><italic>Nerium oleander</italic></td><td align="left" valign="top"> France
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=443.94&link_type=cbs">CBS 443.94</ext-link></td><td align="left" valign="top"><italic>Boeremia exigua</italic> var. <italic>heteromorpha</italic> B
</td><td align="center" valign="top"> GU237935
</td><td align="center" valign="top"> GU237866
</td><td align="center" valign="top"> GU237497
</td><td align="left" valign="top"><italic>Nerium oleander</italic></td><td align="left" valign="top"> Italy
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=569.79&link_type=cbs">CBS 569.79</ext-link>; PD
72/741
</td><td align="left" valign="top"><italic>Boeremia exigua</italic> var. <italic>lilacis</italic> B
</td><td align="center" valign="top"> GU237936
</td><td align="center" valign="top"> GU237892
</td><td align="center" valign="top"> GU237498
</td><td align="left" valign="top"><italic>Syringa vulgaris</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=114.28&link_type=cbs">CBS 114.28</ext-link></td><td align="left" valign="top"><italic>Boeremia exigua</italic> var. <italic>linicola</italic></td><td align="center" valign="top"> GU237937
</td><td align="center" valign="top"> GU237752
</td><td align="center" valign="top"> GU237499
</td><td align="left" valign="top"><italic>Linum usitatissimum</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=116.76&link_type=cbs">CBS 116.76</ext-link>; ATCC
32332; CECT 20022; CECT 20023; IMI 197074
</td><td align="left" valign="top"><italic>Boeremia exigua</italic> var. <italic>linicola</italic> B
</td><td align="center" valign="top"> GU237938
</td><td align="center" valign="top"> GU237754
</td><td align="center" valign="top"> GU237500
</td><td align="left" valign="top"><italic>Linum usitatissimum</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=100167&link_type=cbs">CBS 100167</ext-link>; PD
93/217
</td><td align="left" valign="top"><italic>Boeremia exigua</italic> var. <italic>populi</italic> T
</td><td align="center" valign="top"> GU237939
</td><td align="center" valign="top"> GU237707
</td><td align="center" valign="top"> GU237501
</td><td align="left" valign="top"><italic>Populus</italic> (x) <italic>euramericana</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=101202&link_type=cbs">CBS 101202</ext-link>; PD
82/942
</td><td align="left" valign="top"><italic>Boeremia exigua</italic> var. <italic>populi</italic></td><td align="center" valign="top"> GU237940
</td><td align="center" valign="top"> GU237719
</td><td align="center" valign="top"> GU237502
</td><td align="left" valign="top"><italic>Salix</italic> sp.
</td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=101207&link_type=cbs">CBS 101207</ext-link>; PD
94/614
</td><td align="left" valign="top"><italic>Boeremia exigua</italic> var. <italic>pseudolilacis</italic> T
</td><td align="center" valign="top"> GU237941
</td><td align="center" valign="top"> GU237721
</td><td align="center" valign="top"> GU237503
</td><td align="left" valign="top"><italic>Syringa vulgaris</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=100354&link_type=cbs">CBS 100354</ext-link>; PD
84/448
</td><td align="left" valign="top"><italic>Boeremia exigua</italic> var. <italic>viburni</italic> B
</td><td align="center" valign="top"> GU237944
</td><td align="center" valign="top"> GU237711
</td><td align="center" valign="top"> GU237506
</td><td align="left" valign="top"><italic>Viburnum opulus</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=101211&link_type=cbs">CBS 101211</ext-link>; PD
93/838
</td><td align="left" valign="top"><italic>Boeremia exigua</italic> var. <italic>viburni</italic></td><td align="center" valign="top"> GU237945
</td><td align="center" valign="top"> GU237722
</td><td align="center" valign="top"> GU237507
</td><td align="left" valign="top"><italic>Viburnum</italic> sp.
</td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=109176&link_type=cbs">CBS 109176</ext-link>; CECT
2828; PD 94/1394
</td><td align="left" valign="top"><italic>Boeremia foveata</italic> B
</td><td align="center" valign="top"> GU237946
</td><td align="center" valign="top"> GU237742
</td><td align="center" valign="top"> GU237508
</td><td align="left" valign="top"><italic>Solanum tuberosum</italic></td><td align="left" valign="top"> Bulgaria
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=341.67&link_type=cbs">CBS 341.67</ext-link>; CECT
20055; IMI 331912
</td><td align="left" valign="top"><italic>Boeremia foveata</italic> B
</td><td align="center" valign="top"> GU237947
</td><td align="center" valign="top"> GU237834
</td><td align="center" valign="top"> GU237509
</td><td align="left" valign="top"><italic>Solanum tuberosum</italic></td><td align="left" valign="top"> U.K.
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=366.91&link_type=cbs">CBS 366.91</ext-link>; PD
70/811
</td><td align="left" valign="top"><italic>Boeremia hedericola</italic></td><td align="center" valign="top"> GU237948
</td><td align="center" valign="top"> GU237841
</td><td align="center" valign="top"> GU237510
</td><td align="left" valign="top"><italic>Hedera helix</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=367.91&link_type=cbs">CBS 367.91</ext-link>; PD
87/229
</td><td align="left" valign="top"><italic>Boeremia hedericola</italic> B
</td><td align="center" valign="top"> GU237949
</td><td align="center" valign="top"> GU237842
</td><td align="center" valign="top"> GU237511
</td><td align="left" valign="top"><italic>Hedera helix</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=378.67&link_type=cbs">CBS 378.67</ext-link>; PD
76/276
</td><td align="left" valign="top"><italic>Boeremia lycopersici</italic> B
</td><td align="center" valign="top"> GU237950
</td><td align="center" valign="top"> GU237848
</td><td align="center" valign="top"> GU237512
</td><td align="left" valign="top"><italic>Lycopersicon esculentum</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=109172&link_type=cbs">CBS 109172</ext-link>; PD
84/143
</td><td align="left" valign="top"><italic>Boeremia lycopersici</italic></td><td align="center" valign="top"> GU237951
</td><td align="center" valign="top"> GU237739
</td><td align="center" valign="top"> GU237513
</td><td align="left" valign="top"><italic>Lycopersicon esculentum</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=100353&link_type=cbs">CBS 100353</ext-link>; PD
87/718
</td><td align="left" valign="top"><italic>Boeremia noackiana</italic> B
</td><td align="center" valign="top"> GU237952
</td><td align="center" valign="top"> GU237710
</td><td align="center" valign="top"> GU237514
</td><td align="left" valign="top"><italic>Phaseolus vulgaris</italic></td><td align="left" valign="top"> Guatemala
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=101203&link_type=cbs">CBS 101203</ext-link>; PD
79/1114
</td><td align="left" valign="top"><italic>Boeremia noackiana</italic></td><td align="center" valign="top"> GU237953
</td><td align="center" valign="top"> GU237720
</td><td align="center" valign="top"> GU237515
</td><td align="left" valign="top"><italic>Phaseolus vulgaris</italic></td><td align="left" valign="top"> Colombia
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=109170&link_type=cbs">CBS 109170</ext-link>; PD
75/796
</td><td align="left" valign="top"><italic>Boeremia sambuci-nigrae</italic></td><td align="center" valign="top"> GU237954
</td><td align="center" valign="top"> GU237738
</td><td align="center" valign="top"> GU237516
</td><td align="left" valign="top"><italic>Sambucus nigra</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=629.68&link_type=cbs">CBS 629.68</ext-link>; CECT
20048; IMI 331913; PD 67/753
</td><td align="left" valign="top"><italic>Boeremia sambuci-nigrae</italic> T
</td><td align="center" valign="top"> GU237955
</td><td align="center" valign="top"> GU237897
</td><td align="center" valign="top"> GU237517
</td><td align="left" valign="top"><italic>Sambucus nigra</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=126.93&link_type=cbs">CBS 126.93</ext-link>; PD
73/642
</td><td align="left" valign="top"><italic>Boeremia strasseri</italic></td><td align="center" valign="top"> GU237956
</td><td align="center" valign="top"> GU237773
</td><td align="center" valign="top"> GU237518
</td><td align="left" valign="top"><italic>Mentha</italic> sp.
</td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=261.92&link_type=cbs">CBS 261.92</ext-link>; ATCC
244146; PD 92/318
</td><td align="left" valign="top"><italic>Boeremia strasseri</italic></td><td align="center" valign="top"> GU237957
</td><td align="center" valign="top"> GU237812
</td><td align="center" valign="top"> GU237519
</td><td align="left" valign="top"><italic>Mentha piperita</italic></td><td align="left" valign="top"> U.S.A.
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=109175&link_type=cbs">CBS 109175</ext-link>; PD
79/524
</td><td align="left" valign="top"><italic>Boeremia telephii</italic> B
</td><td align="center" valign="top"> GU237958
</td><td align="center" valign="top"> GU237741
</td><td align="center" valign="top"> GU237520
</td><td align="left" valign="top"><italic>Sedum spectabile</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=760.73&link_type=cbs">CBS 760.73</ext-link>; PD
71/1616
</td><td align="left" valign="top"><italic>Boeremia telephii</italic> B
</td><td align="center" valign="top"> GU237959
</td><td align="center" valign="top"> GU237905
</td><td align="center" valign="top"> GU237521
</td><td align="left" valign="top"><italic>Sedum spectabile</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=148.94&link_type=cbs">CBS 148.94</ext-link></td><td align="left" valign="top"><italic>Chaetasbolisia erysiphoides</italic></td><td align="center" valign="top"><bold>EU754140</bold></td><td align="center" valign="top"> GU237785
</td><td align="center" valign="top"> GU237522
</td><td align="left" valign="top"> Unknown
</td><td align="left" valign="top"> Unknown
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=187.83&link_type=cbs">CBS 187.83</ext-link>; PD
82/128
</td><td align="left" valign="top"><italic>Didymella adianticola</italic> B
</td><td align="center" valign="top"> GU238035
</td><td align="center" valign="top"> GU237796
</td><td align="center" valign="top"> GU237576
</td><td align="left" valign="top"><italic>Polystichum adiantiforme</italic></td><td align="left" valign="top"> U.S.A.
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=258.92&link_type=cbs">CBS 258.92</ext-link>; PD
89/1887
</td><td align="left" valign="top"><italic>Didymella adianticola</italic></td><td align="center" valign="top"> GU238036
</td><td align="center" valign="top"> GU237811
</td><td align="center" valign="top"> GU237577
</td><td align="left" valign="top"><italic>Polystichum adiantiforme</italic></td><td align="left" valign="top"> Costa Rica
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=102634&link_type=cbs">CBS 102634</ext-link>; PD
75/248
</td><td align="left" valign="top"><italic>Didymella applanata</italic></td><td align="center" valign="top"> GU237997
</td><td align="center" valign="top"> GU237726
</td><td align="center" valign="top"> GU237555
</td><td align="left" valign="top"><italic>Rubus idaeus</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=205.63&link_type=cbs">CBS 205.63</ext-link></td><td align="left" valign="top"><italic>Didymella applanata</italic> T
</td><td align="center" valign="top"> GU237998
</td><td align="center" valign="top"> GU237798
</td><td align="center" valign="top"> GU237556
</td><td align="left" valign="top"><italic>Rubus idaeus</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=234.37&link_type=cbs">CBS 234.37</ext-link></td><td align="left" valign="top"><italic>Didymella cannabis</italic></td><td align="center" valign="top"> GU237961
</td><td align="center" valign="top"> GU237804
</td><td align="center" valign="top"> GU237523
</td><td align="left" valign="top"><italic>Cannabis sativa</italic></td><td align="left" valign="top"> Unknown
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=102635&link_type=cbs">CBS 102635</ext-link>; PD
77/1131
</td><td align="left" valign="top"><italic>Didymella catariae</italic></td><td align="center" valign="top"> GU237962
</td><td align="center" valign="top"> GU237727
</td><td align="center" valign="top"> GU237524
</td><td align="left" valign="top"><italic>Nepeta catenaria</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=183.55&link_type=cbs">CBS 183.55</ext-link></td><td align="left" valign="top"><italic>Didymella exigua</italic> T
</td><td align="center" valign="top"><bold>EU754155</bold></td><td align="center" valign="top"> GU237794
</td><td align="center" valign="top"> GU237525
</td><td align="left" valign="top"><italic>Rumex arifolius</italic></td><td align="left" valign="top"> France
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=524.77&link_type=cbs">CBS 524.77</ext-link></td><td align="left" valign="top"><italic>Didymella fabae</italic></td><td align="center" valign="top"> GU237963
</td><td align="center" valign="top"> GU237880
</td><td align="center" valign="top"> GU237526
</td><td align="left" valign="top"><italic>Phaseolus vulgrais</italic></td><td align="left" valign="top"> Belgium
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=649.71&link_type=cbs">CBS 649.71</ext-link></td><td align="left" valign="top"><italic>Didymella fabae</italic></td><td align="center" valign="top"> GU237964
</td><td align="center" valign="top"> GU237902
</td><td align="center" valign="top"> GU237527
</td><td align="left" valign="top"><italic>Vicia faba</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"> PD 83/492
</td><td align="left" valign="top"><italic>Didymella fabae</italic></td><td align="center" valign="top"> GU237965
</td><td align="center" valign="top"> GU237917
</td><td align="center" valign="top"> GU237528
</td><td align="left" valign="top"><italic>Phaseolus vulgaris</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"> PD 84/512
</td><td align="left" valign="top"><italic>Didymella macropodii</italic></td><td align="center" valign="top"> GU237966
</td><td align="center" valign="top"> GU237919
</td><td align="center" valign="top"> GU237529
</td><td align="left" valign="top"> Crucifer
</td><td align="left" valign="top"> Unknown
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=100190&link_type=cbs">CBS 100190</ext-link>; PD
82/736
</td><td align="left" valign="top"><italic>Didymella macropodii</italic></td><td align="center" valign="top"> GU237967
</td><td align="center" valign="top"> GU237708
</td><td align="center" valign="top"> GU237530
</td><td align="left" valign="top"><italic>Brassica napus</italic></td><td align="left" valign="top"> Germany
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=126.54&link_type=cbs">CBS 126.54</ext-link></td><td align="left" valign="top"><italic>Didymella pisi</italic></td><td align="center" valign="top"> GU237968
</td><td align="center" valign="top"> GU237772
</td><td align="center" valign="top"> GU237531
</td><td align="left" valign="top"><italic>Pisum sativum</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=122785&link_type=cbs">CBS 122785</ext-link>; PD
78/517
</td><td align="left" valign="top"><italic>Didymella pisi</italic></td><td align="center" valign="top"> GU237969
</td><td align="center" valign="top"> GU237763
</td><td align="center" valign="top"> GU237532
</td><td align="left" valign="top"><italic>Pisum sativum</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=534.65&link_type=cbs">CBS 534.65</ext-link></td><td align="left" valign="top"><italic>Didymella rabiei</italic></td><td align="center" valign="top"> GU237970
</td><td align="center" valign="top"> GU237886
</td><td align="center" valign="top"> GU237533
</td><td align="left" valign="top"><italic>Cicer arietinum</italic></td><td align="left" valign="top"> India
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=581.83&link_type=cbs">CBS 581.83</ext-link>a
</td><td align="left" valign="top"><italic>Didymella rabiei</italic></td><td align="center" valign="top"> GU237971
</td><td align="center" valign="top"> GU237894
</td><td align="center" valign="top"> GU237534
</td><td align="left" valign="top"><italic>Cicer arietinum</italic></td><td align="left" valign="top"> Syria
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=121.75&link_type=cbs">CBS 121.75</ext-link>; ATCC
32164; IHEM 3403; IMI 194767; PD 73/584
</td><td align="left" valign="top"><italic>Didymella urticicola</italic> T
</td><td align="center" valign="top"> GU237972
</td><td align="center" valign="top"> GU237761
</td><td align="center" valign="top"> GU237535
</td><td align="left" valign="top"><italic>Urtica dioica</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"> PD 73/570
</td><td align="left" valign="top"><italic>Didymella urticicola</italic></td><td align="center" valign="top"> GU237973
</td><td align="center" valign="top"> GU237914
</td><td align="center" valign="top"> GU237536
</td><td align="left" valign="top"><italic>Urtica dioica</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=454.64&link_type=cbs">CBS 454.64</ext-link></td><td align="left" valign="top"><italic>Didymella vitalbina</italic></td><td align="center" valign="top"><bold>FJ515646</bold></td><td align="center" valign="top"><bold>FJ515605</bold></td><td align="center" valign="top"><bold>FJ515623</bold></td><td align="left" valign="top"><italic>Clematis vitalba</italic></td><td align="left" valign="top"> France
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=138.25&link_type=cbs">CBS 138.25</ext-link></td><td align="left" valign="top"><italic>Diplodina coloradensis</italic></td><td align="center" valign="top"><bold>EU754158</bold></td><td align="center" valign="top"> GU237784
</td><td align="center" valign="top"> GU237537
</td><td align="left" valign="top"><italic>Senecio</italic> sp.
</td><td align="left" valign="top"> Unknown
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=172.34&link_type=cbs">CBS 172.34</ext-link></td><td align="left" valign="top"><italic>“Dothiorella ulmi”</italic></td><td align="center" valign="top"><bold>EU754160</bold></td><td align="center" valign="top"> GU237789
</td><td align="center" valign="top"> GU237538
</td><td align="left" valign="top"><italic>Ulmus</italic> sp.
</td><td align="left" valign="top"> U.S.A.
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=125.82&link_type=cbs">CBS 125.82</ext-link>; IMI
1331914; CECT 20044
</td><td align="left" valign="top"><italic>Epicoccum nigrum</italic></td><td align="center" valign="top"> GU237974
</td><td align="center" valign="top"><bold>FJ426995</bold></td><td align="center" valign="top"><bold>FJ427106</bold></td><td align="left" valign="top"> Human
</td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=173.73&link_type=cbs">CBS 173.73</ext-link>; ATCC
24428; IMI 164070
</td><td align="left" valign="top"><italic>Epicoccum nigrum</italic> T
</td><td align="center" valign="top"> GU237975
</td><td align="center" valign="top"><bold>FJ426996</bold></td><td align="center" valign="top"><bold>FJ427107</bold></td><td align="left" valign="top"><italic>Dactylis glomerata</italic></td><td align="left" valign="top"> U.S.A.
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=246.60&link_type=cbs">CBS 246.60</ext-link>; ATCC
22237; ATCC 16652; IMI 081601
</td><td align="left" valign="top"><italic>Epicoccum pimprinum</italic> T
</td><td align="center" valign="top"> GU237976
</td><td align="center" valign="top"><bold>FJ427049</bold></td><td align="center" valign="top"><bold>FJ427159</bold></td><td align="left" valign="top"> Soil
</td><td align="left" valign="top"> India
</td></tr><tr><td align="left" valign="top"> PD 77/1028
</td><td align="left" valign="top"><italic>Epicoccum pimprinum</italic></td><td align="center" valign="top"> GU237977
</td><td align="center" valign="top"><bold>FJ427050</bold></td><td align="center" valign="top"><bold>FJ427160</bold></td><td align="left" valign="top"> Unknown
</td><td align="left" valign="top"> Unknown
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=179.80&link_type=cbs">CBS 179.80</ext-link>; PD
76/1018
</td><td align="left" valign="top"><italic>Epicoccum sorghi</italic></td><td align="center" valign="top"> GU237978
</td><td align="center" valign="top"><bold>FJ427067</bold></td><td align="center" valign="top"><bold>FJ427173</bold></td><td align="left" valign="top"><italic>Sorghum vulgare</italic></td><td align="left" valign="top"> Puerto Rico
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=627.68&link_type=cbs">CBS 627.68</ext-link>; PD
66/926
</td><td align="left" valign="top"><italic>Epicoccum sorghi</italic></td><td align="center" valign="top"> GU237979
</td><td align="center" valign="top"><bold>FJ427072</bold></td><td align="center" valign="top"><bold>FJ427178</bold></td><td align="left" valign="top"><italic>Citrus</italic> sp.
</td><td align="left" valign="top"> France
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=213.55&link_type=cbs">CBS 213.55</ext-link></td><td align="left" valign="top"><italic>Leptosphaerulina americana</italic></td><td align="center" valign="top"> GU237981
</td><td align="center" valign="top"> GU237799
</td><td align="center" valign="top"> GU237539
</td><td align="left" valign="top"><italic>Trifolium pretense</italic></td><td align="left" valign="top"> U.S.A.
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=275.59&link_type=cbs">CBS 275.59</ext-link>; ATCC
13446
</td><td align="left" valign="top"><italic>Leptosphaerulina arachidicola</italic></td><td align="center" valign="top"> GU237983
</td><td align="center" valign="top"> GU237820
</td><td align="center" valign="top"> GU237543
</td><td align="left" valign="top"><italic>Arachis hypochea</italic></td><td align="left" valign="top"> Taiwan
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=317.83&link_type=cbs">CBS 317.83</ext-link></td><td align="left" valign="top"><italic>Leptosphaerulina australis</italic></td><td align="center" valign="top"><bold>EU754166</bold></td><td align="center" valign="top"> GU237829
</td><td align="center" valign="top"> GU237540
</td><td align="left" valign="top"><italic>Eugenia aromatica</italic></td><td align="left" valign="top"> Indonesia
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=939.69&link_type=cbs">CBS 939.69</ext-link></td><td align="left" valign="top"><italic>Leptosphaerulina australis</italic></td><td align="center" valign="top"><bold>EU754167</bold></td><td align="center" valign="top"> GU237911
</td><td align="center" valign="top"> GU237541
</td><td align="left" valign="top"> Soil
</td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=235.58&link_type=cbs">CBS 235.58</ext-link></td><td align="left" valign="top"><italic>Leptosphaerulina trifolii</italic></td><td align="center" valign="top"> GU237982
</td><td align="center" valign="top"> GU237806
</td><td align="center" valign="top"> GU237542
</td><td align="left" valign="top"><italic>Trifolium</italic> sp.
</td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=525.71&link_type=cbs">CBS 525.71</ext-link></td><td align="left" valign="top"><italic>Macroventuria anomochaeta</italic> T
</td><td align="center" valign="top"> GU237984
</td><td align="center" valign="top"> GU237881
</td><td align="center" valign="top"> GU237544
</td><td align="left" valign="top"> decayed canvas
</td><td align="left" valign="top"> South Africa
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=502.72&link_type=cbs">CBS 502.72</ext-link></td><td align="left" valign="top"><italic>Macroventuria anomochaeta</italic></td><td align="center" valign="top"> GU237985
</td><td align="center" valign="top"> GU237873
</td><td align="center" valign="top"> GU237545
</td><td align="left" valign="top"><italic>Medicago sativa</italic></td><td align="left" valign="top"> South Africa
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=526.71&link_type=cbs">CBS 526.71</ext-link></td><td align="left" valign="top"><italic>Macroventuria wentii</italic></td><td align="center" valign="top"> GU237986
</td><td align="center" valign="top"> GU237881
</td><td align="center" valign="top"> GU237546
</td><td align="left" valign="top"> Unidentified plant material
</td><td align="left" valign="top"> U.S.A.
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=432.71&link_type=cbs">CBS 432.71</ext-link></td><td align="left" valign="top"><italic>Microsphaeropsis olivacea</italic></td><td align="center" valign="top"> GU237987
</td><td align="center" valign="top"> GU237863
</td><td align="center" valign="top"> GU237548
</td><td align="left" valign="top"><italic>Sorothamus</italic> sp.
</td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=233.77&link_type=cbs">CBS 233.77</ext-link></td><td align="left" valign="top"><italic>Microsphaeropsis olivacea</italic></td><td align="center" valign="top"> GU237988
</td><td align="center" valign="top"> GU237803
</td><td align="center" valign="top"> GU237549
</td><td align="left" valign="top"><italic>Pinus laricio</italic></td><td align="left" valign="top"> France
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=442.83&link_type=cbs">CBS 442.83</ext-link></td><td align="left" valign="top"><italic>Microsphaeropsis olivacea</italic></td><td align="center" valign="top"><bold>EU754171</bold></td><td align="center" valign="top"> GU237865
</td><td align="center" valign="top"> GU237547
</td><td align="left" valign="top"><italic>Taxus baccata</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=132.96&link_type=cbs">CBS 132.96</ext-link>; PD
93/853
</td><td align="left" valign="top"><italic>Peyronellaea alectorolophi</italic> T
</td><td align="center" valign="top"> GU237989
</td><td align="center" valign="top"> GU237778
</td><td align="center" valign="top"> GU237550
</td><td align="left" valign="top"><italic>Rhinanthus major</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=185.85&link_type=cbs">CBS 185.85</ext-link>; PD
80/1191
</td><td align="left" valign="top"><italic>Peyronellaea americana</italic> B
</td><td align="center" valign="top"> GU237990
</td><td align="center" valign="top"><bold>FJ426972</bold></td><td align="center" valign="top"><bold>FJ427088</bold></td><td align="left" valign="top"><italic>Zea mays</italic></td><td align="left" valign="top"> U.S.A.
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=568.97&link_type=cbs">CBS 568.97</ext-link>; PD
94/1544; ATCC 44494
</td><td align="left" valign="top"><italic>Peyronellaea americana</italic></td><td align="center" valign="top"> GU237991
</td><td align="center" valign="top"><bold>FJ426974</bold></td><td align="center" valign="top"><bold>FJ427090</bold></td><td align="left" valign="top"><italic>Glycine max</italic></td><td align="left" valign="top"> U.S.A.
</td></tr><tr><td align="left" valign="top"> PD 82/1059
</td><td align="left" valign="top"><italic>Peyronellaea americana</italic></td><td align="center" valign="top"> GU237992
</td><td align="center" valign="top"><bold>FJ426980</bold></td><td align="center" valign="top"><bold>FJ427096</bold></td><td align="left" valign="top"> Nematode cyst
</td><td align="left" valign="top"> Unknown
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=360.84&link_type=cbs">CBS 360.84</ext-link></td><td align="left" valign="top"><italic>Peyronellaea anserina</italic> B
</td><td align="center" valign="top"> GU237993
</td><td align="center" valign="top"> GU237839
</td><td align="center" valign="top"> GU237551
</td><td align="left" valign="top"> Potatoflour
</td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=363.91&link_type=cbs">CBS 363.91</ext-link>; PD
79/712
</td><td align="left" valign="top"><italic>Peyronellaea anserina</italic></td><td align="center" valign="top"> GU237994
</td><td align="center" valign="top"> GU237840
</td><td align="center" valign="top"> GU237552
</td><td align="left" valign="top"><italic>Pisum sativum</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=315.90&link_type=cbs">CBS 315.90</ext-link>; PD
80/1190
</td><td align="left" valign="top"><italic>Peyronellaea arachidicola</italic></td><td align="center" valign="top"> GU237995
</td><td align="center" valign="top"> GU237827
</td><td align="center" valign="top"> GU237553
</td><td align="left" valign="top"><italic>Arachis hypogaea</italic></td><td align="left" valign="top"> Zimbabwe
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=333.75&link_type=cbs">CBS 333.75</ext-link>; ATCC
28333; IMI 386092; PREM 44889
</td><td align="left" valign="top"><italic>Peyronellaea arachidicola</italic> T
</td><td align="center" valign="top"> GU237996
</td><td align="center" valign="top"> GU237833
</td><td align="center" valign="top"> GU237554
</td><td align="left" valign="top"><italic>Arachis hypogaea</italic></td><td align="left" valign="top"> South Africa
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=269.93&link_type=cbs">CBS 269.93</ext-link>; PD
78/1087
</td><td align="left" valign="top"><italic>Peyronellaea aurea</italic> B
</td><td align="center" valign="top"> GU237999
</td><td align="center" valign="top"> GU237818
</td><td align="center" valign="top"> GU237557
</td><td align="left" valign="top"><italic>Medicago polymorpha</italic></td><td align="left" valign="top"> New Zealand
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=444.81&link_type=cbs">CBS 444.81</ext-link>; PDDCC
6546
</td><td align="left" valign="top"><italic>Peyronellaea australis</italic> T
</td><td align="center" valign="top"> GU238000
</td><td align="center" valign="top"> GU237867
</td><td align="center" valign="top"> GU237558
</td><td align="left" valign="top"><italic>Actinidia chinensis</italic></td><td align="left" valign="top"> New Zealand
</td></tr><tr><td align="left" valign="top"> PD 77/919
</td><td align="left" valign="top"><italic>Peyronellaea australis</italic></td><td align="center" valign="top"> GU238001
</td><td align="center" valign="top"> GU237915
</td><td align="center" valign="top"> GU237559
</td><td align="left" valign="top"><italic>Actinidea chinensis</italic></td><td align="left" valign="top"> Unknown
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=109.92&link_type=cbs">CBS 109.92</ext-link>; PD
73/1405
</td><td align="left" valign="top"><italic>Peyronellaea calorpreferens</italic> T
</td><td align="center" valign="top"> GU238002
</td><td align="center" valign="top"><bold>FJ426983</bold></td><td align="center" valign="top"><bold>FJ427098</bold></td><td align="left" valign="top"> Undefined food material
</td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=630.97&link_type=cbs">CBS 630.97</ext-link>; ATCC
96683; IMI 361196; PD 96/2022
</td><td align="left" valign="top"><italic>Peyronellaea calorpreferens</italic></td><td align="center" valign="top"> GU238004
</td><td align="center" valign="top"> GU237925
</td><td align="center" valign="top"> GU237560
</td><td align="left" valign="top"><italic>Heterodera glycines</italic></td><td align="left" valign="top"> U.S.A.
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=875.97&link_type=cbs">CBS 875.97</ext-link>; PD
93/1503
</td><td align="left" valign="top"><italic>Peyronellaea calorpreferens</italic></td><td align="center" valign="top"> GU238003
</td><td align="center" valign="top"> GU237908
</td><td align="center" valign="top"> GU237561
</td><td align="left" valign="top"> Indoor environment
</td><td align="left" valign="top"> U.S.A.
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123380&link_type=cbs">CBS 123380</ext-link>; PD
84/1013
</td><td align="left" valign="top"><italic>Peyronellaea coffeae-arabicae</italic> T
</td><td align="center" valign="top"> GU238005
</td><td align="center" valign="top"><bold>FJ426993</bold></td><td align="center" valign="top"><bold>FJ427104</bold></td><td align="left" valign="top"><italic>Coffea arabica</italic></td><td align="left" valign="top"> Ethiopia
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123398&link_type=cbs">CBS 123398</ext-link>; PD
84/1014
</td><td align="left" valign="top"><italic>Peyronellaea coffeae-arabicae</italic></td><td align="center" valign="top"> GU238006
</td><td align="center" valign="top"><bold>FJ426994</bold></td><td align="center" valign="top"><bold>FJ427105</bold></td><td align="left" valign="top"><italic>Coffea arabica</italic></td><td align="left" valign="top"> Ethiopia
</td></tr><tr><td align="left" valign="top"> PD 92/1460
</td><td align="left" valign="top"><italic>Peyronellaea curtisii</italic></td><td align="center" valign="top"> GU238012
</td><td align="center" valign="top"><bold>FJ427041</bold></td><td align="center" valign="top"><bold>FJ427151</bold></td><td align="left" valign="top"><italic>Sprekelia</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=251.92&link_type=cbs">CBS 251.92</ext-link>; PD
86/1145
</td><td align="left" valign="top"><italic>Peyronellaea curtisii</italic> B
</td><td align="center" valign="top"> GU238013
</td><td align="center" valign="top"><bold>FJ427038</bold></td><td align="center" valign="top"><bold>FJ427148</bold></td><td align="left" valign="top"><italic>Nerine</italic> sp.
</td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=377.91&link_type=cbs">CBS 377.91</ext-link>; PD
79/210
</td><td align="left" valign="top"><italic>Peyronellaea eucalyptica</italic> B
</td><td align="center" valign="top"> GU238007
</td><td align="center" valign="top"> GU237846
</td><td align="center" valign="top"> GU237562
</td><td align="left" valign="top"><italic>Eucalyptus</italic> sp.
</td><td align="left" valign="top"> Australia
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=508.91&link_type=cbs">CBS 508.91</ext-link>; PD
73/1413
</td><td align="left" valign="top"><italic>Peyronellaea eucalyptica</italic></td><td align="center" valign="top"> GU238008
</td><td align="center" valign="top"> GU237878
</td><td align="center" valign="top"> GU237563
</td><td align="left" valign="top"> Water
</td><td align="left" valign="top"> Croatia
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=302.79&link_type=cbs">CBS 302.79</ext-link>; PD
79/1156
</td><td align="left" valign="top"><italic>Peyronellaea gardeniae</italic></td><td align="center" valign="top"><bold>GQ387596</bold></td><td align="center" valign="top"><bold>FJ427002</bold></td><td align="center" valign="top"><bold>FJ427113</bold></td><td align="left" valign="top"> Air
</td><td align="left" valign="top"> Netherlands Antilles
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=626.68&link_type=cbs">CBS 626.68</ext-link>; IMI
108771
</td><td align="left" valign="top"><italic>Peyronellaea gardeniae</italic> T
</td><td align="center" valign="top"><bold>GQ387595</bold></td><td align="center" valign="top"><bold>FJ427003</bold></td><td align="center" valign="top"><bold>FJ427114</bold></td><td align="left" valign="top"><italic>Gardenia jasminoides</italic></td><td align="left" valign="top"> India
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=464.97&link_type=cbs">CBS 464.97</ext-link>; MUCL
9882
</td><td align="left" valign="top"><italic>Peyronellaea glomerata</italic></td><td align="center" valign="top"> GU238009
</td><td align="center" valign="top"><bold>FJ427012</bold></td><td align="center" valign="top"><bold>FJ427123</bold></td><td align="left" valign="top"> Indoor environment
</td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=528.66&link_type=cbs">CBS 528.66</ext-link>; PD
63/590
</td><td align="left" valign="top"><italic>Peyronellaea glomerata</italic> B
</td><td align="center" valign="top"><bold>EU754184</bold></td><td align="center" valign="top"><bold>FJ427013</bold></td><td align="center" valign="top"><bold>FJ427124</bold></td><td align="left" valign="top"><italic>Chrysanthemum</italic> sp.
</td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=103.25&link_type=cbs">CBS 103.25</ext-link></td><td align="left" valign="top"><italic>Peyronellaea lethalis</italic></td><td align="center" valign="top"> GU238010
</td><td align="center" valign="top"> GU237729
</td><td align="center" valign="top"> GU237564
</td><td align="left" valign="top"> Unknown
</td><td align="left" valign="top"> Unknown
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=463.69&link_type=cbs">CBS 463.69</ext-link></td><td align="left" valign="top"><italic>Peyronellaea musae</italic> B
</td><td align="center" valign="top"> GU238011
</td><td align="center" valign="top"><bold>FJ427026</bold></td><td align="center" valign="top"><bold>FJ427136</bold></td><td align="left" valign="top"><italic>Mangifera indica</italic></td><td align="left" valign="top"> India
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=377.93&link_type=cbs">CBS 377.93</ext-link>; PD
80/976
</td><td align="left" valign="top"><italic>Peyronellaea obtusa</italic> B
</td><td align="center" valign="top"> GU238014
</td><td align="center" valign="top"> GU237847
</td><td align="center" valign="top"> GU237565
</td><td align="left" valign="top"><italic>Daucus carota</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=391.93&link_type=cbs">CBS 391.93</ext-link>; PD
80/87
</td><td align="left" valign="top"><italic>Peyronellaea obtusa</italic> B
</td><td align="center" valign="top"> GU238015
</td><td align="center" valign="top"> GU237858
</td><td align="center" valign="top"> GU237566
</td><td align="left" valign="top"><italic>Spinacia oleracea</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=318.90&link_type=cbs">CBS 318.90</ext-link>; PD
81/729
</td><td align="left" valign="top"><italic>Peyronellaea pinodella</italic></td><td align="center" valign="top"> GU238016
</td><td align="center" valign="top"><bold>FJ427051</bold></td><td align="center" valign="top"><bold>FJ427161</bold></td><td align="left" valign="top"><italic>Pisum sativum</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=531.66&link_type=cbs">CBS 531.66</ext-link></td><td align="left" valign="top"><italic>Peyronellaea pinodella</italic> B
</td><td align="center" valign="top"> GU238017
</td><td align="center" valign="top"><bold>FJ427052</bold></td><td align="center" valign="top"><bold>FJ427162</bold></td><td align="left" valign="top"><italic>Trifolium pratense</italic></td><td align="left" valign="top"> U.S.A.
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=100580&link_type=cbs">CBS 100580</ext-link>; PD
98/1135
</td><td align="left" valign="top"><italic>Peyronellaea pinodella</italic></td><td align="center" valign="top"> GU238018
</td><td align="center" valign="top"> GU237713
</td><td align="center" valign="top"> GU237567
</td><td align="left" valign="top"><italic>Glycine max</italic></td><td align="left" valign="top"> Hungary
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=567.97&link_type=cbs">CBS 567.97</ext-link>; PD
97/2160
</td><td align="left" valign="top"><italic>Peyronellaea pinodella</italic></td><td align="center" valign="top"> GU238019
</td><td align="center" valign="top"> GU237891
</td><td align="center" valign="top"> GU237568
</td><td align="left" valign="top"><italic>Glycine max</italic></td><td align="left" valign="top"> Hungary
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=159.78&link_type=cbs">CBS 159.78</ext-link>b
</td><td align="left" valign="top"><italic>Peyronellaea pinodes</italic></td><td align="center" valign="top"> GU238020
</td><td align="center" valign="top"> GU237786
</td><td align="center" valign="top"> GU237569
</td><td align="left" valign="top"><italic>Pisum sativum</italic></td><td align="left" valign="top"> Iraq
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=285.49&link_type=cbs">CBS 285.49</ext-link></td><td align="left" valign="top"><italic>Peyronellaea pinodes</italic></td><td align="center" valign="top"> GU238022
</td><td align="center" valign="top"> GU237823
</td><td align="center" valign="top"> GU237571
</td><td align="left" valign="top"><italic>Primula auricula</italic></td><td align="left" valign="top"> Switzerland
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=235.55&link_type=cbs">CBS 235.55</ext-link></td><td align="left" valign="top"><italic>Peyronellaea pinodes</italic></td><td align="center" valign="top"> GU238021
</td><td align="center" valign="top"> GU237805
</td><td align="center" valign="top"> GU237570
</td><td align="left" valign="top"> Unknown
</td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=525.77&link_type=cbs">CBS 525.77</ext-link></td><td align="left" valign="top"><italic>Peyronellaea pinodes</italic></td><td align="center" valign="top"> GU238023
</td><td align="center" valign="top"> GU237883
</td><td align="center" valign="top"> GU237572
</td><td align="left" valign="top"><italic>Pisum sativum</italic></td><td align="left" valign="top"> Belgium
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=525.77&link_type=cbs">CBS 525.77</ext-link>a
</td><td align="left" valign="top"><italic>Peyronellaea pinodes</italic></td><td align="center" valign="top"> GU238024
</td><td align="center" valign="top"> GU237882
</td><td align="center" valign="top"> GU237573
</td><td align="left" valign="top"><italic>Pisum sativum</italic></td><td align="left" valign="top"> Belgium
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=539.66&link_type=cbs">CBS 539.66</ext-link>; ATCC
16791; IMI 122266; PD 64/914
</td><td align="left" valign="top"><italic>Peyronellaea pomorum</italic> var. <italic>pomorum</italic> B
</td><td align="center" valign="top"> GU238028
</td><td align="center" valign="top"><bold>FJ427056</bold></td><td align="center" valign="top"><bold>FJ427166</bold></td><td align="left" valign="top"><italic>Polygonum tataricum</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=285.76&link_type=cbs">CBS 285.76</ext-link>; ATCC
26241; IMI 176742; VKM F-1843
</td><td align="left" valign="top"><italic>Peyronellaea pomorum</italic> var. <italic>circinata</italic> T
</td><td align="center" valign="top"> GU238025
</td><td align="center" valign="top"><bold>FJ427053</bold></td><td align="center" valign="top"><bold>FJ427163</bold></td><td align="left" valign="top"><italic>Heracleum dissectum</italic></td><td align="left" valign="top"> Russia
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=286.76&link_type=cbs">CBS 286.76</ext-link>; ATCC
26242; IMI 176743; VKM F-1844
</td><td align="left" valign="top"><italic>Peyronellaea pomorum</italic> var. <italic>circinata</italic></td><td align="center" valign="top"> GU238026
</td><td align="center" valign="top"><bold>FJ427054</bold></td><td align="center" valign="top"><bold>FJ427164</bold></td><td align="left" valign="top"><italic>Allium nutans</italic></td><td align="left" valign="top"> Russia
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=388.80&link_type=cbs">CBS 388.80</ext-link>; PREM
45736
</td><td align="left" valign="top"><italic>Peyronellaea pomorum</italic> var. <italic>cyanea</italic> T
</td><td align="center" valign="top"> GU238027
</td><td align="center" valign="top"><bold>FJ427055</bold></td><td align="center" valign="top"><bold>FJ427165</bold></td><td align="left" valign="top"><italic>Triticum</italic> sp.
</td><td align="left" valign="top"> South Africa
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=381.96&link_type=cbs">CBS 381.96</ext-link>; PD
71/706
</td><td align="left" valign="top"><italic>Peyronellaea protuberans</italic> B
</td><td align="center" valign="top"> GU238029
</td><td align="center" valign="top"> GU237853
</td><td align="center" valign="top"> GU237574
</td><td align="left" valign="top"><italic>Lycium halifolium</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=281.83&link_type=cbs">CBS 281.83</ext-link></td><td align="left" valign="top"><italic>Peyronellaea sancta</italic> T
</td><td align="center" valign="top"> GU238030
</td><td align="center" valign="top"><bold>FJ427063</bold></td><td align="center" valign="top"><bold>FJ427170</bold></td><td align="left" valign="top"><italic>Ailanthus altissima</italic></td><td align="left" valign="top"> South Africa
</td></tr><tr><td align="left" valign="top"> LEV 15292
</td><td align="left" valign="top"><italic>Peyronellaea sancta</italic></td><td align="center" valign="top"> GU238031
</td><td align="center" valign="top"><bold>FJ427065</bold></td><td align="center" valign="top"><bold>FJ427172</bold></td><td align="left" valign="top"><italic>Gleditsia triacantha</italic></td><td align="left" valign="top"> Unknown
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=110.92&link_type=cbs">CBS 110.92</ext-link>; PD
76/1010
</td><td align="left" valign="top"><italic>Peyronellaea subglomerata</italic> B
</td><td align="center" valign="top"> GU238032
</td><td align="center" valign="top"><bold>FJ427080</bold></td><td align="center" valign="top"><bold>FJ427186</bold></td><td align="left" valign="top"><italic>Triticum</italic> sp.
</td><td align="left" valign="top"> U.S.A.
</td></tr><tr><td align="left" valign="top"> PD 78/1090
</td><td align="left" valign="top"><italic>Peyronellaea subglomerata</italic></td><td align="center" valign="top"> GU238033
</td><td align="center" valign="top"><bold>FJ427081</bold></td><td align="center" valign="top"><bold>FJ427187</bold></td><td align="left" valign="top"><italic>Zea mays</italic></td><td align="left" valign="top"> Unknown
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=588.69&link_type=cbs">CBS 588.69</ext-link></td><td align="left" valign="top"><italic>Peyronellaea zeae-maydis</italic> T
</td><td align="center" valign="top"><bold>EU754186</bold></td><td align="center" valign="top"><bold>FJ427086</bold></td><td align="center" valign="top"><bold>FJ427190</bold></td><td align="left" valign="top"><italic>Zea mays</italic></td><td align="left" valign="top"> U.S.A.
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=179.97&link_type=cbs">CBS 179.97</ext-link></td><td align="left" valign="top"><italic>Phoma acetosellae</italic></td><td align="center" valign="top"> GU238034
</td><td align="center" valign="top"> GU237793
</td><td align="center" valign="top"> GU237575
</td><td align="left" valign="top"><italic>Rumex hydrolapathum</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=379.93&link_type=cbs">CBS 379.93</ext-link>; PD
82/945
</td><td align="left" valign="top"><italic>Phoma aliena</italic></td><td align="center" valign="top"> GU238037
</td><td align="center" valign="top"> GU237851
</td><td align="center" valign="top"> GU237578
</td><td align="left" valign="top"><italic>Berberis</italic> sp.
</td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=877.97&link_type=cbs">CBS 877.97</ext-link>; PD
94/1401
</td><td align="left" valign="top"><italic>Phoma aliena</italic></td><td align="center" valign="top"> GU238038
</td><td align="center" valign="top"> GU237910
</td><td align="center" valign="top"> GU237579
</td><td align="left" valign="top"><italic>Buxus sempervirens</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=381.91&link_type=cbs">CBS 381.91</ext-link>; PD
79/1110
</td><td align="left" valign="top"><italic>Phoma anigozanthi</italic> B
</td><td align="center" valign="top"> GU238039
</td><td align="center" valign="top"> GU237852
</td><td align="center" valign="top"> GU237580
</td><td align="left" valign="top"><italic>Anigozanthus maugleisii</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=107.96&link_type=cbs">CBS 107.96</ext-link>; PD
73/598
</td><td align="left" valign="top"><italic>Phoma aquilegiicola</italic> B
</td><td align="center" valign="top"> GU238041
</td><td align="center" valign="top"> GU237735
</td><td align="center" valign="top"> GU237582
</td><td align="left" valign="top"><italic>Aconitum pyramidale</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=108.96&link_type=cbs">CBS 108.96</ext-link>; PD
79/611
</td><td align="left" valign="top"><italic>Phoma aquilegiicola</italic> B
</td><td align="center" valign="top"> GU238042
</td><td align="center" valign="top"> GU237736
</td><td align="center" valign="top"> GU237583
</td><td align="left" valign="top"><italic>Aquilegia</italic> sp.
</td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=125.93&link_type=cbs">CBS 125.93</ext-link>; PD
77/1029
</td><td align="left" valign="top"><italic>Phoma arachidis-hypogaeae</italic> B
</td><td align="center" valign="top"> GU238043
</td><td align="center" valign="top"> GU237771
</td><td align="center" valign="top"> GU237584
</td><td align="left" valign="top"><italic>Arachis hypogaea</italic></td><td align="left" valign="top"> India
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=383.67&link_type=cbs">CBS 383.67</ext-link>; PD
65/223
</td><td align="left" valign="top"><italic>Phoma aubrietiae</italic> B
</td><td align="center" valign="top"> GU238044
</td><td align="center" valign="top"> GU237854
</td><td align="center" valign="top"> GU237585
</td><td align="left" valign="top"><italic>Aubrietia hybrida</italic> cv. Superbissima
</td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=627.97&link_type=cbs">CBS 627.97</ext-link>; PD
70/714
</td><td align="left" valign="top"><italic>Phoma aubrietiae</italic> B
</td><td align="center" valign="top"> GU238045
</td><td align="center" valign="top"> GU237895
</td><td align="center" valign="top"> GU237586
</td><td align="left" valign="top"><italic>Aubrietia</italic> sp.
</td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=714.85&link_type=cbs">CBS 714.85</ext-link>; PD
74/265
</td><td align="left" valign="top"><italic>Phoma bellidis</italic> B
</td><td align="center" valign="top"> GU238046
</td><td align="center" valign="top"> GU237904
</td><td align="center" valign="top"> GU237587
</td><td align="left" valign="top"><italic>Bellis perennis</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"> PD 94/886
</td><td align="left" valign="top"><italic>Phoma bellidis</italic></td><td align="center" valign="top"> GU238047
</td><td align="center" valign="top"> GU237923
</td><td align="center" valign="top"> GU237581
</td><td align="left" valign="top"><italic>Bellis</italic> sp.
</td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=109942&link_type=cbs">CBS 109942</ext-link>; PD
84/402
</td><td align="left" valign="top"><italic>Phoma boeremae</italic> T
</td><td align="center" valign="top"> GU238048
</td><td align="center" valign="top"><bold>FJ426982</bold></td><td align="center" valign="top"><bold>FJ427097</bold></td><td align="left" valign="top"><italic>Medicago littoralis</italic> cv. Harbinger
</td><td align="left" valign="top"> Australia
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=120105&link_type=cbs">CBS 120105</ext-link></td><td align="left" valign="top"><italic>Phoma brasiliensis</italic> T
</td><td align="center" valign="top"> GU238049
</td><td align="center" valign="top"> GU237760
</td><td align="center" valign="top"> GU237588
</td><td align="left" valign="top"><italic>Amaranthus</italic> sp.
</td><td align="left" valign="top"> Brazil
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=357.84&link_type=cbs">CBS 357.84</ext-link></td><td align="left" valign="top"><italic>Phoma bulgarica</italic> T
</td><td align="center" valign="top"> GU238050
</td><td align="center" valign="top"> GU237837
</td><td align="center" valign="top"> GU237589
</td><td align="left" valign="top"><italic>Trachystemon orientale</italic></td><td align="left" valign="top"> Bulgaria
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=124515&link_type=cbs">CBS 124515</ext-link>; PD
82/1058
</td><td align="left" valign="top"><italic>Phoma bulgarica</italic></td><td align="center" valign="top"> GU238051
</td><td align="center" valign="top"> GU237768
</td><td align="center" valign="top"> GU237590
</td><td align="left" valign="top"><italic>Trachystemon orientale</italic></td><td align="left" valign="top"> Bulgaria
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=448.83&link_type=cbs">CBS 448.83</ext-link></td><td align="left" valign="top"><italic>Phoma calidophila</italic> T
</td><td align="center" valign="top"> GU238052
</td><td align="center" valign="top"><bold>FJ427059</bold></td><td align="center" valign="top"><bold>FJ427168</bold></td><td align="left" valign="top"> Soil
</td><td align="left" valign="top"> Egypt
</td></tr><tr><td align="left" valign="top"> PD 84/109
</td><td align="left" valign="top"><italic>Phoma calidophila</italic></td><td align="center" valign="top"> GU238053
</td><td align="center" valign="top"><bold>FJ427060</bold></td><td align="center" valign="top"><bold>FJ427169</bold></td><td align="left" valign="top"><italic>Cucumis sativus</italic></td><td align="left" valign="top"> Europe
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=128.93&link_type=cbs">CBS 128.93</ext-link>; PD
79/140
</td><td align="left" valign="top"><italic>Phoma chenopodiicola</italic> B
</td><td align="center" valign="top"> GU238055
</td><td align="center" valign="top"> GU237775
</td><td align="center" valign="top"> GU237591
</td><td align="left" valign="top"><italic>Chenopodium quinoa</italic> cv. Sajana
</td><td align="left" valign="top"> Peru
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=129.93&link_type=cbs">CBS 129.93</ext-link>; PD
89/803
</td><td align="left" valign="top"><italic>Phoma chenopodiicola</italic></td><td align="center" valign="top"> GU238056
</td><td align="center" valign="top"> GU237776
</td><td align="center" valign="top"> GU237592
</td><td align="left" valign="top"><italic>Chenopodium quinoa</italic> cv. Sajana
</td><td align="left" valign="top"> Peru
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=102.66&link_type=cbs">CBS 102.66</ext-link></td><td align="left" valign="top"><italic>Phoma clematidina</italic></td><td align="center" valign="top"><bold>FJ515630</bold></td><td align="center" valign="top"><bold>FJ426988</bold></td><td align="center" valign="top"><bold>FJ427099</bold></td><td align="left" valign="top"><italic>Clematis</italic> sp.
</td><td align="left" valign="top"> U.K.
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=108.79&link_type=cbs">CBS 108.79</ext-link>; PD
78/522
</td><td align="left" valign="top"><italic>Phoma clematidina</italic> T
</td><td align="center" valign="top"><bold>FJ515632</bold></td><td align="center" valign="top"><bold>FJ426989</bold></td><td align="center" valign="top"><bold>FJ427100</bold></td><td align="left" valign="top"><italic>Clematis</italic> sp.
</td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=507.63&link_type=cbs">CBS 507.63</ext-link>; MUCL
9574; PD 07/03486747
</td><td align="left" valign="top"><italic>Phoma clematidis-rectae</italic> T
</td><td align="center" valign="top"><bold>FJ515647</bold></td><td align="center" valign="top"><bold>FJ515606</bold></td><td align="center" valign="top"><bold>FJ515624</bold></td><td align="left" valign="top"><italic>Clematis</italic> sp.
</td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"> PD 95/1958
</td><td align="left" valign="top"><italic>Phoma clematidis-rectae</italic></td><td align="center" valign="top"><bold>FJ515648</bold></td><td align="center" valign="top"><bold>FJ515607</bold></td><td align="center" valign="top"><bold>FJ515625</bold></td><td align="left" valign="top"><italic>Clematis</italic> sp.
</td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=100409&link_type=cbs">CBS 100409</ext-link></td><td align="left" valign="top"><italic>Phoma commelinicicola</italic> B
</td><td align="center" valign="top"> GU238057
</td><td align="center" valign="top"> GU237712
</td><td align="center" valign="top"> GU237593
</td><td align="left" valign="top"><italic>Tradescantia</italic> sp.
</td><td align="left" valign="top"> New Zealand
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=100311&link_type=cbs">CBS 100311</ext-link></td><td align="left" valign="top"><italic>Phoma complanata</italic></td><td align="center" valign="top"><bold>EU754181</bold></td><td align="center" valign="top"> GU237709
</td><td align="center" valign="top"> GU237594
</td><td align="left" valign="top"><italic>Heracleum sphondylium</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=268.92&link_type=cbs">CBS 268.92</ext-link>; PD 75/3
</td><td align="left" valign="top"><italic>Phoma complanata</italic></td><td align="center" valign="top"><bold>EU754180</bold></td><td align="center" valign="top"> GU237815
</td><td align="center" valign="top"> GU237595
</td><td align="left" valign="top"><italic>Angelica sylvestris</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=506.91&link_type=cbs">CBS 506.91</ext-link>; IMI
215229; PD 91/876
</td><td align="left" valign="top"><italic>Phoma costarricensis</italic> B
</td><td align="center" valign="top"> GU238058
</td><td align="center" valign="top"> GU237876
</td><td align="center" valign="top"> GU237596
</td><td align="left" valign="top"><italic>Coffea</italic> sp.
</td><td align="left" valign="top"> Nicaragua
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=497.91&link_type=cbs">CBS 497.91</ext-link>; PD
79/209
</td><td align="left" valign="top"><italic>Phoma costarricensis</italic></td><td align="center" valign="top"> GU238059
</td><td align="center" valign="top"> GU237870
</td><td align="center" valign="top"> GU237597
</td><td align="left" valign="top"><italic>Coffea arabica</italic></td><td align="left" valign="top"> Unknown
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=193.82&link_type=cbs">CBS 193.82</ext-link></td><td align="left" valign="top"><italic>Phoma crystallifera</italic> T
</td><td align="center" valign="top"> GU238060
</td><td align="center" valign="top"> GU237797
</td><td align="center" valign="top"> GU237598
</td><td align="left" valign="top"><italic>Chamaespartium sagittale</italic></td><td align="left" valign="top"> Austria
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=124513&link_type=cbs">CBS 124513</ext-link>; PD
73/1414
</td><td align="left" valign="top"><italic>Phoma dactylidis</italic> T
</td><td align="center" valign="top"> GU238061
</td><td align="center" valign="top"> GU237766
</td><td align="center" valign="top"> GU237599
</td><td align="left" valign="top"><italic>Dactylis glomerata</italic></td><td align="left" valign="top"> U.S.A.
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=133.93&link_type=cbs">CBS 133.93</ext-link>; PD
88/961; IMI 173142
</td><td align="left" valign="top"><italic>Phoma destructiva</italic> var. <italic>destructiva</italic></td><td align="center" valign="top"> GU238064
</td><td align="center" valign="top"> GU237779
</td><td align="center" valign="top"> GU237602
</td><td align="left" valign="top"><italic>Solanum lycopersicum</italic></td><td align="left" valign="top"> Guadeloupe
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=378.73&link_type=cbs">CBS 378.73</ext-link>; CECT
2877
</td><td align="left" valign="top"><italic>Phoma destructiva</italic> var. <italic>destructiva</italic> B
</td><td align="center" valign="top"> GU238063
</td><td align="center" valign="top"> GU237849
</td><td align="center" valign="top"> GU237601
</td><td align="left" valign="top"><italic>Lycopersicon esculentum</italic></td><td align="left" valign="top"> Tonga
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=162.78&link_type=cbs">CBS 162.78</ext-link>; PD
77/725
</td><td align="left" valign="top"><italic>Phoma destructiva</italic> var. <italic>diversispora</italic></td><td align="center" valign="top"> GU238062
</td><td align="center" valign="top"> GU237788
</td><td align="center" valign="top"> GU237600
</td><td align="left" valign="top"><italic>Lycopersicon esculentum</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=507.91&link_type=cbs">CBS 507.91</ext-link>; PD
74/148
</td><td align="left" valign="top"><italic>Phoma dictamnicola</italic> B
</td><td align="center" valign="top"> GU238065
</td><td align="center" valign="top"> GU237877
</td><td align="center" valign="top"> GU237603
</td><td align="left" valign="top"><italic>Dictamnus albus</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=109179&link_type=cbs">CBS 109179</ext-link>; PD
90/835-1
</td><td align="left" valign="top"><italic>Phoma digitalis</italic></td><td align="center" valign="top"> GU238066
</td><td align="center" valign="top"> GU237744
</td><td align="center" valign="top"> GU237604
</td><td align="left" valign="top"><italic>Digitalis</italic> sp.
</td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=229.79&link_type=cbs">CBS 229.79</ext-link>; LEV
7660
</td><td align="left" valign="top"><italic>Phoma digitalis</italic> B
</td><td align="center" valign="top"> GU238067
</td><td align="center" valign="top"> GU237802
</td><td align="center" valign="top"> GU237605
</td><td align="left" valign="top"><italic>Digitalis purpurea</italic></td><td align="left" valign="top"> New Zealand
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=346.82&link_type=cbs">CBS 346.82</ext-link></td><td align="left" valign="top"><italic>Phoma dimorpha</italic> T
</td><td align="center" valign="top"> GU238068
</td><td align="center" valign="top"> GU237835
</td><td align="center" valign="top"> GU237606
</td><td align="left" valign="top"><italic>Opuntiae</italic> sp.
</td><td align="left" valign="top"> Spain
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=186.83&link_type=cbs">CBS 186.83</ext-link>; PD
82/47
</td><td align="left" valign="top"><italic>Phoma draconis</italic> B
</td><td align="center" valign="top"> GU238070
</td><td align="center" valign="top"> GU237795
</td><td align="center" valign="top"> GU237607
</td><td align="left" valign="top"><italic>Dracaena</italic> sp.
</td><td align="left" valign="top"> Rwanda
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123.93&link_type=cbs">CBS 123.93</ext-link>; PD
77/1148
</td><td align="left" valign="top"><italic>Phoma eupatorii</italic> B
</td><td align="center" valign="top"> GU238071
</td><td align="center" valign="top"> GU237764
</td><td align="center" valign="top"> GU237608
</td><td align="left" valign="top"><italic>Eupatorium cannabinum</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=374.91&link_type=cbs">CBS 374.91</ext-link>; PD
78/391
</td><td align="left" valign="top"><italic>Phoma eupyrena</italic> B
</td><td align="center" valign="top"> GU238072
</td><td align="center" valign="top"><bold>FJ426999</bold></td><td align="center" valign="top"><bold>FJ427110</bold></td><td align="left" valign="top"><italic>Solanum tuberosum</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=527.66&link_type=cbs">CBS 527.66</ext-link>; ATCC
22238
</td><td align="left" valign="top"><italic>Phoma eupyrena</italic> B
</td><td align="center" valign="top"> GU238073
</td><td align="center" valign="top"><bold>FJ427000</bold></td><td align="center" valign="top"><bold>FJ427111</bold></td><td align="left" valign="top"> Soil
</td><td align="left" valign="top"> Germany
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=633.92&link_type=cbs">CBS 633.92</ext-link>; ATCC
36786; VKM MF-325
</td><td align="left" valign="top"><italic>Phoma fungicola</italic></td><td align="center" valign="top"><bold>EU754127</bold></td><td align="center" valign="top"> GU237900
</td><td align="center" valign="top"> GU237609
</td><td align="left" valign="top"><italic>Microsphaera alphitoides</italic> on <italic>Quercus</italic> sp.
</td><td align="left" valign="top"> Ukraine
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=112.96&link_type=cbs">CBS 112.96</ext-link></td><td align="left" valign="top"><italic>Phoma glaucii</italic></td><td align="center" valign="top"> GU238077
</td><td align="center" valign="top"> GU237750
</td><td align="center" valign="top"> GU237610
</td><td align="left" valign="top"><italic>Dicentra</italic> sp.
</td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=114.96&link_type=cbs">CBS 114.96</ext-link>; PD
94/888
</td><td align="left" valign="top"><italic>Phoma glaucii</italic> B
</td><td align="center" valign="top"><bold>FJ515649</bold></td><td align="center" valign="top"><bold>FJ515609</bold></td><td align="center" valign="top"><bold>FJ515627</bold></td><td align="left" valign="top"><italic>Chelidonium majus</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=377.67&link_type=cbs">CBS 377.67</ext-link></td><td align="left" valign="top"><italic>Phoma gossypiicola</italic> B
</td><td align="center" valign="top"> GU238079
</td><td align="center" valign="top"> GU237845
</td><td align="center" valign="top"> GU237611
</td><td align="left" valign="top"><italic>Gossypium hirsutum</italic></td><td align="left" valign="top"> U.S.A.
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=104.80&link_type=cbs">CBS 104.80</ext-link>; PD
74/1017
</td><td align="left" valign="top"><italic>Phoma henningsii</italic> B
</td><td align="center" valign="top"> GU238081
</td><td align="center" valign="top"> GU237731
</td><td align="center" valign="top"> GU237612
</td><td align="left" valign="top"><italic>Acacia mearnesii</italic></td><td align="left" valign="top"> Kenya
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=502.91&link_type=cbs">CBS 502.91</ext-link>; PD
86/276
</td><td align="left" valign="top"><italic>Phoma herbarum</italic></td><td align="center" valign="top"> GU238082
</td><td align="center" valign="top"> GU237874
</td><td align="center" valign="top"> GU237613
</td><td align="left" valign="top"><italic>Nerium</italic> sp.
</td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=615.75&link_type=cbs">CBS 615.75</ext-link>; PD
73/665; IMI 199779
</td><td align="left" valign="top"><italic>Phoma herbarum</italic> B
</td><td align="center" valign="top"><bold>EU880896</bold></td><td align="center" valign="top"><bold>FJ427022</bold></td><td align="center" valign="top"><bold>FJ427133</bold></td><td align="left" valign="top"><italic>Rosa multiflora</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=629.97&link_type=cbs">CBS 629.97</ext-link>; PD
76/1017
</td><td align="left" valign="top"><italic>Phoma herbicola</italic> B
</td><td align="center" valign="top"> GU238083
</td><td align="center" valign="top"> GU237898
</td><td align="center" valign="top"> GU237614
</td><td align="left" valign="top"> Water
</td><td align="left" valign="top"> U.S.A.
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=105.80&link_type=cbs">CBS 105.80</ext-link>; PD
75/908
</td><td align="left" valign="top"><italic>Phoma huancayensis</italic> T
</td><td align="center" valign="top"> GU238084
</td><td align="center" valign="top"> GU237732
</td><td align="center" valign="top"> GU237615
</td><td align="left" valign="top"><italic>Solanum</italic> sp.
</td><td align="left" valign="top"> Peru
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=390.93&link_type=cbs">CBS 390.93</ext-link>; PD
77/1173
</td><td align="left" valign="top"><italic>Phoma huancayensis</italic></td><td align="center" valign="top"> GU238085
</td><td align="center" valign="top"> GU237857
</td><td align="center" valign="top"> GU237616
</td><td align="left" valign="top"><italic>Chenopodium quinoa</italic></td><td align="left" valign="top"> Peru
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=220.85&link_type=cbs">CBS 220.85</ext-link></td><td align="left" valign="top"><italic>Phoma humicola</italic> B
</td><td align="center" valign="top"> GU238086
</td><td align="center" valign="top"> GU237800
</td><td align="center" valign="top"> GU237617
</td><td align="left" valign="top"><italic>Franseria</italic> sp.
</td><td align="left" valign="top"> U.S.A.
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123394&link_type=cbs">CBS 123394</ext-link></td><td align="left" valign="top"><italic>Phoma infossa</italic></td><td align="center" valign="top"> GU238088
</td><td align="center" valign="top"><bold>FJ427024</bold></td><td align="center" valign="top"><bold>FJ427134</bold></td><td align="left" valign="top"><italic>Fraxinus pennsylvanica</italic></td><td align="left" valign="top"> Argentina
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123395&link_type=cbs">CBS 123395</ext-link></td><td align="left" valign="top"><italic>Phoma infossa</italic> T
</td><td align="center" valign="top"> GU238089
</td><td align="center" valign="top"><bold>FJ427025</bold></td><td align="center" valign="top"><bold>FJ427135</bold></td><td align="left" valign="top"><italic>Fraxinus pennsylvanica</italic></td><td align="left" valign="top"> Argentina
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=252.92&link_type=cbs">CBS 252.92</ext-link>; PD
80/1144
</td><td align="left" valign="top"><italic>Phoma insulana</italic> B
</td><td align="center" valign="top"> GU238090
</td><td align="center" valign="top"> GU237810
</td><td align="center" valign="top"> GU237618
</td><td align="left" valign="top"><italic>Olea europaea</italic></td><td align="left" valign="top"> Greece
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=124.93&link_type=cbs">CBS 124.93</ext-link>; PD
87/269
</td><td align="left" valign="top"><italic>Phoma labilis</italic> B
</td><td align="center" valign="top"> GU238091
</td><td align="center" valign="top"> GU237765
</td><td align="center" valign="top"> GU237619
</td><td align="left" valign="top"><italic>Solanum lycopersicum</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=479.93&link_type=cbs">CBS 479.93</ext-link>; PD
70/93
</td><td align="left" valign="top"><italic>Phoma labilis</italic></td><td align="center" valign="top"> GU238092
</td><td align="center" valign="top"> GU237868
</td><td align="center" valign="top"> GU237620
</td><td align="left" valign="top"><italic>Rosa</italic> sp.
</td><td align="left" valign="top"> Israel
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=347.82&link_type=cbs">CBS 347.82</ext-link></td><td align="left" valign="top"><italic>Phoma longicolla</italic></td><td align="center" valign="top"> GU238094
</td><td align="center" valign="top"> GU237836
</td><td align="center" valign="top"> GU237621
</td><td align="left" valign="top"><italic>Opuntiae</italic> sp.
</td><td align="left" valign="top"> Spain
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=124514&link_type=cbs">CBS 124514</ext-link>; PD
80/1189; VPRI 1239
</td><td align="left" valign="top"><italic>Phoma longicolla</italic> T
</td><td align="center" valign="top"> GU238095
</td><td align="center" valign="top"> GU237767
</td><td align="center" valign="top"> GU237622
</td><td align="left" valign="top"><italic>Opuntiae</italic> sp.
</td><td align="left" valign="top"> Spain
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=223.69&link_type=cbs">CBS 223.69</ext-link></td><td align="left" valign="top"><italic>Phoma macrostoma</italic> var. <italic>incolorata</italic> B
</td><td align="center" valign="top"> GU238096
</td><td align="center" valign="top"> GU237801
</td><td align="center" valign="top"> GU237623
</td><td align="left" valign="top"><italic>Acer pseudoplatanus</italic></td><td align="left" valign="top"> Switzerland
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=109173&link_type=cbs">CBS 109173</ext-link>; PD
83/908
</td><td align="left" valign="top"><italic>Phoma macrostoma</italic> var. <italic>incolorata</italic> B
</td><td align="center" valign="top"> GU238097
</td><td align="center" valign="top"> GU237740
</td><td align="center" valign="top"> GU237624
</td><td align="left" valign="top"><italic>Malus sylvestris</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=529.66&link_type=cbs">CBS 529.66</ext-link>; PD
66/521
</td><td align="left" valign="top"><italic>Phoma macrostoma</italic> var. <italic>macrostoma</italic> B
</td><td align="center" valign="top"> GU238098
</td><td align="center" valign="top"> GU237885
</td><td align="center" valign="top"> GU237625
</td><td align="left" valign="top"><italic>Malus sylvestris</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=482.95&link_type=cbs">CBS 482.95</ext-link></td><td align="left" valign="top"><italic>Phoma macrostoma</italic> var. <italic>macrostoma</italic></td><td align="center" valign="top"> GU238099
</td><td align="center" valign="top"> GU237869
</td><td align="center" valign="top"> GU237626
</td><td align="left" valign="top"><italic>Larix decidua</italic></td><td align="left" valign="top"> Germany
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=259.92&link_type=cbs">CBS 259.92</ext-link>; IMI
286996; PD 91/272
</td><td align="left" valign="top"><italic>Phoma matteuciicola</italic> B
</td><td align="center" valign="top"> GU238100
</td><td align="center" valign="top"> GU237812
</td><td align="center" valign="top"> GU237627
</td><td align="left" valign="top"><italic>Matteuccia struthiopteris</italic></td><td align="left" valign="top"> Canada
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=112.53&link_type=cbs">CBS 112.53</ext-link></td><td align="left" valign="top"><italic>Phoma medicaginis</italic> var. <italic>macrospora</italic> B
</td><td align="center" valign="top"> GU238101
</td><td align="center" valign="top"> GU237749
</td><td align="center" valign="top"> GU237628
</td><td align="left" valign="top"><italic>Medicago sativa</italic></td><td align="left" valign="top"> U.S.A.
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=404.65&link_type=cbs">CBS 404.65</ext-link>; IMI
116999
</td><td align="left" valign="top"><italic>Phoma medicaginis</italic> var. <italic>macrospora</italic> B
</td><td align="center" valign="top"> GU238102
</td><td align="center" valign="top"> GU237859
</td><td align="center" valign="top"> GU237629
</td><td align="left" valign="top"><italic>Medicago sativa</italic></td><td align="left" valign="top"> Canada
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=316.90&link_type=cbs">CBS 316.90</ext-link></td><td align="left" valign="top"><italic>Phoma medicaginis</italic> var. <italic>medicaginis</italic></td><td align="center" valign="top"> GU238103
</td><td align="center" valign="top"> GU237828
</td><td align="center" valign="top"> GU237630
</td><td align="left" valign="top"><italic>Medicago sativa</italic></td><td align="left" valign="top"> Czech Republic
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=105.95&link_type=cbs">CBS 105.95</ext-link></td><td align="left" valign="top"><italic>Phoma microchlamydospora</italic> T
</td><td align="center" valign="top"> GU238104
</td><td align="center" valign="top"><bold>FJ427028</bold></td><td align="center" valign="top"><bold>FJ427138</bold></td><td align="left" valign="top"><italic>Eucalyptus</italic> sp.
</td><td align="left" valign="top"> U.K.
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=491.90&link_type=cbs">CBS 491.90</ext-link></td><td align="left" valign="top"><italic>Phoma microchlamydospora</italic></td><td align="center" valign="top"> GU238105
</td><td align="center" valign="top"><bold>FJ427029</bold></td><td align="center" valign="top"><bold>FJ427139</bold></td><td align="left" valign="top"> Unidentified vegetable
</td><td align="left" valign="top"> U.K.
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=315.83&link_type=cbs">CBS 315.83</ext-link></td><td align="left" valign="top"><italic>Phoma minor</italic></td><td align="center" valign="top"> GU238106
</td><td align="center" valign="top"> GU237826
</td><td align="center" valign="top"> GU237631
</td><td align="left" valign="top"><italic>Syzygium aromaticum</italic></td><td align="left" valign="top"> Indonesia
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=325.82&link_type=cbs">CBS 325.82</ext-link></td><td align="left" valign="top"><italic>Phoma minor</italic> T
</td><td align="center" valign="top"> GU238107
</td><td align="center" valign="top"> GU237831
</td><td align="center" valign="top"> GU237632
</td><td align="left" valign="top"><italic>Syzygium aromaticum</italic></td><td align="left" valign="top"> Indonesia
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=110.79&link_type=cbs">CBS 110.79</ext-link>; PD
65/8875; MUCL 8247
</td><td align="left" valign="top"><italic>Phoma multirostrata</italic></td><td align="center" valign="top"> GU238110
</td><td align="center" valign="top"><bold>FJ427030</bold></td><td align="center" valign="top"><bold>FJ427140</bold></td><td align="left" valign="top"><italic>Cucumis sativus</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=274.60&link_type=cbs">CBS 274.60</ext-link>; IMI
081598
</td><td align="left" valign="top"><italic>Phoma multirostrata</italic> T
</td><td align="center" valign="top"> GU238111
</td><td align="center" valign="top"><bold>FJ427031</bold></td><td align="center" valign="top"><bold>FJ427141</bold></td><td align="left" valign="top"> Soil
</td><td align="left" valign="top"> India
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=368.65&link_type=cbs">CBS 368.65</ext-link>; PD
92/1757; HACC 154
</td><td align="left" valign="top"><italic>Phoma multirostrata</italic></td><td align="center" valign="top"> GU238112
</td><td align="center" valign="top"><bold>FJ427033</bold></td><td align="center" valign="top"><bold>FJ427143</bold></td><td align="left" valign="top"> Soil
</td><td align="left" valign="top"> India
</td></tr><tr><td align="left" valign="top"> PD 83/48
</td><td align="left" valign="top"><italic>Phoma multirostrata</italic></td><td align="center" valign="top"> GU238113
</td><td align="center" valign="top"><bold>FJ427037</bold></td><td align="center" valign="top"><bold>FJ427147</bold></td><td align="left" valign="top"><italic>Cucumis sativus</italic></td><td align="left" valign="top"> Unknown
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=117.93&link_type=cbs">CBS 117.93</ext-link>; PD
83/90
</td><td align="left" valign="top"><italic>Phoma nebulosa</italic></td><td align="center" valign="top"> GU238114
</td><td align="center" valign="top"> GU237757
</td><td align="center" valign="top"> GU237633
</td><td align="left" valign="top"><italic>Mercurialis perennis</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=503.75&link_type=cbs">CBS 503.75</ext-link>; ATCC
32163; DSM 63391; IMI 194766; PD 75/4
</td><td align="left" valign="top"><italic>Phoma nebulosa</italic> B
</td><td align="center" valign="top"> GU238115
</td><td align="center" valign="top"> GU237875
</td><td align="center" valign="top"> GU237634
</td><td align="left" valign="top"><italic>Urtica dioica</italic></td><td align="left" valign="top"> Austria
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=358.71&link_type=cbs">CBS 358.71</ext-link></td><td align="left" valign="top"><italic>Phoma negriana</italic> B
</td><td align="center" valign="top"> GU238116
</td><td align="center" valign="top"> GU237838
</td><td align="center" valign="top"> GU237635
</td><td align="left" valign="top"><italic>Vitis vinifera</italic></td><td align="left" valign="top"> Germany
</td></tr><tr><td align="left" valign="top"> PD 79/74
</td><td align="left" valign="top"><italic>Phoma negriana</italic></td><td align="center" valign="top"> GU238117
</td><td align="center" valign="top"> GU237916
</td><td align="center" valign="top"> GU237636
</td><td align="left" valign="top"><italic>Vitis vinifera</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=116.96&link_type=cbs">CBS 116.96</ext-link>; PD
95/7930
</td><td align="left" valign="top"><italic>Phoma nigripycnidia</italic> B
</td><td align="center" valign="top"> GU238118
</td><td align="center" valign="top"> GU237756
</td><td align="center" valign="top"> GU237637
</td><td align="left" valign="top"><italic>Vicia cracca</italic></td><td align="left" valign="top"> Russia
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=114.93&link_type=cbs">CBS 114.93</ext-link>; PD
74/228
</td><td align="left" valign="top"><italic>Phoma novae- verbascicola</italic></td><td align="center" valign="top"> GU238119
</td><td align="center" valign="top"> GU237753
</td><td align="center" valign="top"> GU237638
</td><td align="left" valign="top"><italic>Verbascum</italic> sp.
</td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=127.93&link_type=cbs">CBS 127.93</ext-link>; PD
92/347
</td><td align="left" valign="top"><italic>Phoma novae-verbascicola</italic> B
</td><td align="center" valign="top"> GU238120
</td><td align="center" valign="top"> GU237774
</td><td align="center" valign="top"> GU237639
</td><td align="left" valign="top"><italic>Verbascum densiflorum</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=654.77&link_type=cbs">CBS 654.77</ext-link></td><td align="left" valign="top"><italic>Phoma omnivirens</italic></td><td align="center" valign="top"> GU238122
</td><td align="center" valign="top"><bold>FJ427043</bold></td><td align="center" valign="top"><bold>FJ427153</bold></td><td align="left" valign="top"> Unknown
</td><td align="left" valign="top"> India
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=991.95&link_type=cbs">CBS 991.95</ext-link></td><td align="left" valign="top"><italic>Phoma omnivirens</italic></td><td align="center" valign="top"> GU238121
</td><td align="center" valign="top"><bold>FJ427044</bold></td><td align="center" valign="top"><bold>FJ427154</bold></td><td align="left" valign="top"> Soil
</td><td align="left" valign="top"> Papua New Guinea
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=560.81&link_type=cbs">CBS 560.81</ext-link>; PD
92/1569; PDDCC 6614
</td><td align="left" valign="top"><italic>Phoma paspali</italic> T
</td><td align="center" valign="top"> GU238124
</td><td align="center" valign="top"><bold>FJ427048</bold></td><td align="center" valign="top"><bold>FJ427158</bold></td><td align="left" valign="top"><italic>Paspalum dilatatum</italic></td><td align="left" valign="top"> New Zealand
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=561.81&link_type=cbs">CBS 561.81</ext-link>; PDDCC
6615
</td><td align="left" valign="top"><italic>Phoma paspali</italic></td><td align="center" valign="top"> GU238125
</td><td align="center" valign="top"> GU237889
</td><td align="center" valign="top"> GU237640
</td><td align="left" valign="top"><italic>Lolium perenne</italic></td><td align="left" valign="top"> New Zealand
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=124516&link_type=cbs">CBS 124516</ext-link>; PD
84/453
</td><td align="left" valign="top"><italic>Phoma pedeiae</italic></td><td align="center" valign="top"> GU238126
</td><td align="center" valign="top"> GU237769
</td><td align="center" valign="top"> GU237641
</td><td align="left" valign="top"><italic>Orchidaceae</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=124517&link_type=cbs">CBS 124517</ext-link>; PD
92/612A
</td><td align="left" valign="top"><italic>Phoma pedeiae</italic> T
</td><td align="center" valign="top"> GU238127
</td><td align="center" valign="top"> GU237770
</td><td align="center" valign="top"> GU237642
</td><td align="left" valign="top"><italic>Schefflera elegantissima</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=267.92&link_type=cbs">CBS 267.92</ext-link>; PD
76/1014
</td><td align="left" valign="top"><italic>Phoma pereupyrena</italic> T
</td><td align="center" valign="top"> GU238128
</td><td align="center" valign="top"> GU237814
</td><td align="center" valign="top"> GU237643
</td><td align="left" valign="top"><italic>Coffea arabica</italic></td><td align="left" valign="top"> India
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=268.93&link_type=cbs">CBS 268.93</ext-link>;
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=108.93&link_type=cbs">CBS 108.93</ext-link>; PD
88/720
</td><td align="left" valign="top"><italic>Phoma piperis</italic> B
</td><td align="center" valign="top"> GU238129
</td><td align="center" valign="top"> GU237816
</td><td align="center" valign="top"> GU237644
</td><td align="left" valign="top"><italic>Peperomia pereskifolia</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"> PD 90/2011
</td><td align="left" valign="top"><italic>Phoma piperis</italic></td><td align="center" valign="top"> GU238130
</td><td align="center" valign="top"> GU237921
</td><td align="center" valign="top"> GU237645
</td><td align="left" valign="top"><italic>Peperomia</italic> sp.
</td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=284.93&link_type=cbs">CBS 284.93</ext-link>; PD
75/907
</td><td align="left" valign="top"><italic>Phoma plurivora</italic></td><td align="center" valign="top"> GU238131
</td><td align="center" valign="top"> GU237822
</td><td align="center" valign="top"> GU237646
</td><td align="left" valign="top"><italic>Medicago sativa</italic></td><td align="left" valign="top"> Australia
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=558.81&link_type=cbs">CBS 558.81</ext-link>; PDDCC
6873
</td><td align="left" valign="top"><italic>Phoma plurivora</italic> T
</td><td align="center" valign="top"> GU238132
</td><td align="center" valign="top"> GU237888
</td><td align="center" valign="top"> GU237647
</td><td align="left" valign="top"><italic>Setaria</italic> sp.
</td><td align="left" valign="top"> New Zealand
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=109181&link_type=cbs">CBS 109181</ext-link>; PD
83/757
</td><td align="left" valign="top"><italic>Phoma polemonii</italic> B
</td><td align="center" valign="top"> GU238133
</td><td align="center" valign="top"> GU237746
</td><td align="center" valign="top"> GU237648
</td><td align="left" valign="top"><italic>Polemonium caeruleum</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=116.93&link_type=cbs">CBS 116.93</ext-link>; PD
71/884
</td><td align="left" valign="top"><italic>Phoma poolensis</italic> B
</td><td align="center" valign="top"> GU238134
</td><td align="center" valign="top"> GU237755
</td><td align="center" valign="top"> GU237649
</td><td align="left" valign="top"><italic>Antirrhinum majus</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=113.20&link_type=cbs">CBS 113.20</ext-link>; PD
92/774
</td><td align="left" valign="top"><italic>Phoma poolensis</italic></td><td align="center" valign="top"> GU238135
</td><td align="center" valign="top"> GU237751
</td><td align="center" valign="top"> GU237650
</td><td align="left" valign="top"> Unknown
</td><td align="left" valign="top"> Unknown
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=372.91&link_type=cbs">CBS 372.91</ext-link>; PD
75/690
</td><td align="left" valign="top"><italic>Phoma putaminum</italic> B
</td><td align="center" valign="top"> GU238137
</td><td align="center" valign="top"> GU237843
</td><td align="center" valign="top"> GU237651
</td><td align="left" valign="top"><italic>Ulmus</italic> sp.
</td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=130.69&link_type=cbs">CBS 130.69</ext-link>; CECT
20054; IMI 331916
</td><td align="left" valign="top"><italic>Phoma putaminum</italic> B
</td><td align="center" valign="top"> GU238138
</td><td align="center" valign="top"> GU237777
</td><td align="center" valign="top"> GU237652
</td><td align="left" valign="top"><italic>Malus sylvestris</italic></td><td align="left" valign="top"> Denmark
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=109177&link_type=cbs">CBS 109177</ext-link>; LEV
15165; PD 2000/9941
</td><td align="left" valign="top"><italic>Phoma rhei</italic> B
</td><td align="center" valign="top"> GU238139
</td><td align="center" valign="top"> GU237743
</td><td align="center" valign="top"> GU237653
</td><td align="left" valign="top"><italic>Rheum rhaponticum</italic></td><td align="left" valign="top"> New Zealand
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=298.89&link_type=cbs">CBS 298.89</ext-link></td><td align="left" valign="top"><italic>Phoma saxea</italic></td><td align="center" valign="top"> GU238140
</td><td align="center" valign="top"> GU237824
</td><td align="center" valign="top"> GU237654
</td><td align="left" valign="top"> Limestone
</td><td align="left" valign="top"> Germany
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=419.92&link_type=cbs">CBS 419.92</ext-link></td><td align="left" valign="top"><italic>Phoma saxea</italic> T
</td><td align="center" valign="top"> GU238141
</td><td align="center" valign="top"> GU237860
</td><td align="center" valign="top"> GU237655
</td><td align="left" valign="top"> Corroded mediterranean marble
</td><td align="left" valign="top"> Germany
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=122.93&link_type=cbs">CBS 122.93</ext-link>; PD
77/1049
</td><td align="left" valign="top"><italic>Phoma selaginellicola</italic> B
</td><td align="center" valign="top"> GU238142
</td><td align="center" valign="top"> GU237762
</td><td align="center" valign="top"> GU237656
</td><td align="left" valign="top"><italic>Selaginella</italic> sp.
</td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=160.78&link_type=cbs">CBS 160.78</ext-link>; LEV
11451
</td><td align="left" valign="top"><italic>Phoma senecionis</italic> B
</td><td align="center" valign="top"> GU238143
</td><td align="center" valign="top"> GU237787
</td><td align="center" valign="top"> GU237657
</td><td align="left" valign="top"><italic>Senecio jacobaea</italic></td><td align="left" valign="top"> New Zealand
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=249.92&link_type=cbs">CBS 249.92</ext-link>; PD
78/1088
</td><td align="left" valign="top"><italic>Phoma subherbarum</italic></td><td align="center" valign="top"> GU238144
</td><td align="center" valign="top"> GU237808
</td><td align="center" valign="top"> GU237658
</td><td align="left" valign="top"><italic>Solanum</italic> sp.
</td><td align="left" valign="top"> Peru
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=250.92&link_type=cbs">CBS 250.92</ext-link>; DAOM
171914; PD 92/371
</td><td align="left" valign="top"><italic>Phoma subherbarum</italic> B
</td><td align="center" valign="top"> GU238145
</td><td align="center" valign="top"> GU237809
</td><td align="center" valign="top"> GU237659
</td><td align="left" valign="top"><italic>Solanum</italic> sp.
</td><td align="left" valign="top"> Peru
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=305.79&link_type=cbs">CBS 305.79</ext-link>A; DAOM
170848
</td><td align="left" valign="top"><italic>Phoma subherbarum</italic></td><td align="center" valign="top"> GU238146
</td><td align="center" valign="top"> GU237825
</td><td align="center" valign="top"> GU237660
</td><td align="left" valign="top"><italic>Zea mays</italic></td><td align="left" valign="top"> Peru
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=135.93&link_type=cbs">CBS 135.93</ext-link>; PD
83/87
</td><td align="left" valign="top"><italic>Phoma sylvatica</italic> B
</td><td align="center" valign="top"> GU238147
</td><td align="center" valign="top"> GU237781
</td><td align="center" valign="top"> GU237661
</td><td align="left" valign="top"><italic>Melampyrum pratense</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=874.97&link_type=cbs">CBS 874.97</ext-link>; PD
93/764
</td><td align="left" valign="top"><italic>Phoma sylvatica</italic> B
</td><td align="center" valign="top"> GU238148
</td><td align="center" valign="top"> GU237907
</td><td align="center" valign="top"> GU237662
</td><td align="left" valign="top"><italic>Melampyrum pratense</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=436.75&link_type=cbs">CBS 436.75</ext-link></td><td align="left" valign="top"><italic>Phoma tropica</italic> T
</td><td align="center" valign="top"> GU238149
</td><td align="center" valign="top"> GU237864
</td><td align="center" valign="top"> GU237663
</td><td align="left" valign="top"><italic>Saintpaulia ionantha</italic></td><td align="left" valign="top"> Germany
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=876.97&link_type=cbs">CBS 876.97</ext-link>; PD
82/1008
</td><td align="left" valign="top"><italic>Phoma versabilis</italic> B
</td><td align="center" valign="top"> GU238152
</td><td align="center" valign="top"> GU237909
</td><td align="center" valign="top"> GU237664
</td><td align="left" valign="top"><italic>Silene</italic> sp.
</td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"> PD 2000/1379
</td><td align="left" valign="top"><italic>Phoma versabilis</italic></td><td align="center" valign="top"> GU238153
</td><td align="center" valign="top"> GU237913
</td><td align="center" valign="top"> GU237665
</td><td align="left" valign="top"><italic>Stellaria media</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=500.91&link_type=cbs">CBS 500.91</ext-link>; PD
83/322
</td><td align="left" valign="top"><italic>Phoma viburnicola</italic> B
</td><td align="center" valign="top"> GU238154
</td><td align="center" valign="top"> GU237871
</td><td align="center" valign="top"> GU237666
</td><td align="left" valign="top"><italic>Ilex aquifolium</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=523.73&link_type=cbs">CBS 523.73</ext-link>; PD
69/800
</td><td align="left" valign="top"><italic>Phoma viburnicola</italic> B
</td><td align="center" valign="top"> GU238155
</td><td align="center" valign="top"> GU237879
</td><td align="center" valign="top"> GU237667
</td><td align="left" valign="top"><italic>Viburnum cassioides</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=383.68&link_type=cbs">CBS 383.68</ext-link></td><td align="left" valign="top"><italic>Phoma xanthina</italic> B
</td><td align="center" valign="top"> GU238157
</td><td align="center" valign="top"> GU237855
</td><td align="center" valign="top"> GU237668
</td><td align="left" valign="top"><italic>Delphinium</italic> sp.
</td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"> PD 84/407
</td><td align="left" valign="top"><italic>Phoma xanthina</italic></td><td align="center" valign="top"> GU238158
</td><td align="center" valign="top"> GU237918
</td><td align="center" valign="top"> GU237669
</td><td align="left" valign="top"><italic>Delphinium</italic> sp.
</td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=131.93&link_type=cbs">CBS 131.93</ext-link>; PD
69/140
</td><td align="left" valign="top"><italic>Phoma zantedeschiae</italic></td><td align="center" valign="top"> GU238159
</td><td align="center" valign="top"><bold>FJ427084</bold></td><td align="center" valign="top"><bold>FJ427188</bold></td><td align="left" valign="top"><italic>Calla</italic> sp.
</td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=105.96&link_type=cbs">CBS 105.96</ext-link>; PD
74/230
</td><td align="left" valign="top"><italic>Stagonosporopsis actaeae</italic> B
</td><td align="center" valign="top"> GU238165
</td><td align="center" valign="top"> GU237733
</td><td align="center" valign="top"> GU237670
</td><td align="left" valign="top"><italic>Cimicifuga simplex</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=106.96&link_type=cbs">CBS 106.96</ext-link>; PD
94/1318
</td><td align="left" valign="top"><italic>Stagonosporopsis actaeae</italic> T
</td><td align="center" valign="top"> GU238166
</td><td align="center" valign="top"> GU237734
</td><td align="center" valign="top"> GU237671
</td><td align="left" valign="top"><italic>Actaea spicata</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=176.93&link_type=cbs">CBS 176.93</ext-link>; PD
86/547
</td><td align="left" valign="top"><italic>Stagonosporopsis ajacis</italic></td><td align="center" valign="top"> GU238167
</td><td align="center" valign="top"> GU237790
</td><td align="center" valign="top"> GU237672
</td><td align="left" valign="top"><italic>Delphinium</italic> sp.
</td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=177.93&link_type=cbs">CBS 177.93</ext-link>; PD
90/115
</td><td align="left" valign="top"><italic>Stagonosporopsis ajacis</italic> T
</td><td align="center" valign="top"> GU238168
</td><td align="center" valign="top"> GU237791
</td><td align="center" valign="top"> GU237673
</td><td align="left" valign="top"><italic>Delphinium</italic> sp.
</td><td align="left" valign="top"> Kenya
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=101.80&link_type=cbs">CBS 101.80</ext-link>; PD
75/909; IMI 386090
</td><td align="left" valign="top"><italic>Stagonosporopsis andigena</italic> B
</td><td align="center" valign="top"> GU238169
</td><td align="center" valign="top"> GU237714
</td><td align="center" valign="top"> GU237674
</td><td align="left" valign="top"><italic>Solanum</italic> sp.
</td><td align="left" valign="top"> Peru
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=269.80&link_type=cbs">CBS 269.80</ext-link>; PD
75/914
</td><td align="left" valign="top"><italic>Stagonosporopsis andigena</italic></td><td align="center" valign="top"> GU238170
</td><td align="center" valign="top"> GU237817
</td><td align="center" valign="top"> GU237675
</td><td align="left" valign="top"><italic>Solanum</italic> sp.
</td><td align="left" valign="top"> Peru
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=102636&link_type=cbs">CBS 102636</ext-link>; PD
73/1409
</td><td align="left" valign="top"><italic>Stagonosporopsis artemisiicola</italic> B
</td><td align="center" valign="top"> GU238171
</td><td align="center" valign="top"> GU237728
</td><td align="center" valign="top"> GU237676
</td><td align="left" valign="top"><italic>Artemisia dracunculus</italic></td><td align="left" valign="top"> France
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=178.25&link_type=cbs">CBS 178.25</ext-link>; MUCL
9915
</td><td align="left" valign="top"><italic>Stagonosporopsis astragali</italic> B
</td><td align="center" valign="top"> GU238172
</td><td align="center" valign="top"> GU237792
</td><td align="center" valign="top"> GU237677
</td><td align="left" valign="top"><italic>Astragalus</italic> sp.
</td><td align="left" valign="top"> Unknown
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=248.90&link_type=cbs">CBS 248.90</ext-link></td><td align="left" valign="top"><italic>Stagonosporopsis caricae</italic></td><td align="center" valign="top"> GU238175
</td><td align="center" valign="top"> GU237807
</td><td align="center" valign="top"> GU237680
</td><td align="left" valign="top"><italic>Carica papaya</italic></td><td align="left" valign="top"> Chile
</td></tr><tr><td align="left" valign="top"> PD 06/03082531
</td><td align="left" valign="top"><italic>Stagonosporopsis caricae</italic></td><td align="center" valign="top"> GU238176
</td><td align="center" valign="top"> GU237912
</td><td align="center" valign="top"> GU237681
</td><td align="left" valign="top"><italic>Carica papaya</italic></td><td align="left" valign="top"> Brazil
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=282.76&link_type=cbs">CBS 282.76</ext-link></td><td align="left" valign="top"><italic>Stagonosporopsis caricae</italic></td><td align="center" valign="top"> GU238177
</td><td align="center" valign="top"> GU237821
</td><td align="center" valign="top"> GU237682
</td><td align="left" valign="top"><italic>Brassica</italic> sp.
</td><td align="left" valign="top"> Indonesia
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=713.85&link_type=cbs">CBS 713.85</ext-link>; ATCC
76027; PD 83/826
</td><td align="left" valign="top"><italic>Stagonosporopsis crystalliniformis</italic> T
</td><td align="center" valign="top"> GU238178
</td><td align="center" valign="top"> GU237903
</td><td align="center" valign="top"> GU237683
</td><td align="left" valign="top"><italic>Lycopersicon esculentum</italic></td><td align="left" valign="top"> Colombia
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=771.85&link_type=cbs">CBS 771.85</ext-link>; IMI
386091; PD 85/772
</td><td align="left" valign="top"><italic>Stagonosporopsis crystalliniformis</italic></td><td align="center" valign="top"> GU238179
</td><td align="center" valign="top"> GU237906
</td><td align="center" valign="top"> GU237684
</td><td align="left" valign="top"><italic>Solanum tuberosum</italic></td><td align="left" valign="top"> Colombia
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=109171&link_type=cbs">CBS 109171</ext-link>; PD
91/310; PDDCC 272
</td><td align="left" valign="top"><italic>Stagonosporopsis cucurbitacearum</italic></td><td align="center" valign="top"> GU238180
</td><td align="center" valign="top"> GU237922
</td><td align="center" valign="top"> GU237685
</td><td align="left" valign="top"><italic>Cucurbita</italic> sp.
</td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=133.96&link_type=cbs">CBS 133.96</ext-link>; PD
79/127
</td><td align="left" valign="top"><italic>Stagonosporopsis cucurbitacearum</italic></td><td align="center" valign="top"> GU238181
</td><td align="center" valign="top"> GU237780
</td><td align="center" valign="top"> GU237686
</td><td align="left" valign="top"><italic>Cucurbita</italic> sp.
</td><td align="left" valign="top"> New Zealand
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=631.68&link_type=cbs">CBS 631.68</ext-link>; PD
68/147
</td><td align="left" valign="top"><italic>Stagonosporopsis dennisii</italic> B
</td><td align="center" valign="top"> GU238182
</td><td align="center" valign="top"> GU237899
</td><td align="center" valign="top"> GU237687
</td><td align="left" valign="top"><italic>Solidago floribunda</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=135.96&link_type=cbs">CBS 135.96</ext-link>; IMI
19337; PD 95/4756
</td><td align="left" valign="top"><italic>Stagonosporopsis dennisii</italic></td><td align="center" valign="top"> GU238183
</td><td align="center" valign="top"> GU237782
</td><td align="center" valign="top"> GU237688
</td><td align="left" valign="top"><italic>Solidago canadensis</italic></td><td align="left" valign="top"> Canada
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=320.90&link_type=cbs">CBS 320.90</ext-link>; PD
86/932
</td><td align="left" valign="top"><italic>Stagonosporopsis dorenboschii</italic> B
</td><td align="center" valign="top"> GU238184
</td><td align="center" valign="top"> GU237830
</td><td align="center" valign="top"> GU237689
</td><td align="left" valign="top"><italic>Physostegia virginiana</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=426.90&link_type=cbs">CBS 426.90</ext-link>; IMI
386093; PD 86/551
</td><td align="left" valign="top"><italic>Stagonosporopsis dorenboschii</italic> T
</td><td align="center" valign="top"> GU238185
</td><td align="center" valign="top"> GU237862
</td><td align="center" valign="top"> GU237690
</td><td align="left" valign="top"><italic>Physostegia virginiana</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=109182&link_type=cbs">CBS 109182</ext-link>; PD
74/231
</td><td align="left" valign="top"><italic>Stagonosporopsis heliopsidis</italic> B
</td><td align="center" valign="top"> GU238186
</td><td align="center" valign="top"> GU237747
</td><td align="center" valign="top"> GU237691
</td><td align="left" valign="top"><italic>Heliopsis patula</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"> PD 95/6189; DAOM 221138
</td><td align="left" valign="top"><italic>Stagonosporopsis heliopsidis</italic></td><td align="center" valign="top"> GU238187
</td><td align="center" valign="top"> GU237924
</td><td align="center" valign="top"> GU237692
</td><td align="left" valign="top"><italic>Ambrosia artemisiifolia</italic></td><td align="left" valign="top"> Canada
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=104.42&link_type=cbs">CBS 104.42</ext-link></td><td align="left" valign="top"><italic>Stagonosporopsis hortensis</italic> B
</td><td align="center" valign="top"> GU238198
</td><td align="center" valign="top"> GU237730
</td><td align="center" valign="top"> GU237703
</td><td align="left" valign="top"> Unknown
</td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=572.85&link_type=cbs">CBS 572.85</ext-link>; PD
79/269
</td><td align="left" valign="top"><italic>Stagonosporopsis hortensis</italic> B
</td><td align="center" valign="top"> GU238199
</td><td align="center" valign="top"> GU237893
</td><td align="center" valign="top"> GU237704
</td><td align="left" valign="top"><italic>Phaseolus vulgaris</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=425.90&link_type=cbs">CBS 425.90</ext-link>; PD
81/520
</td><td align="left" valign="top"><italic>Stagonosporopsis ligulicola</italic> var. <italic>inoxydabilis</italic> T
</td><td align="center" valign="top"> GU238188
</td><td align="center" valign="top"> GU237861
</td><td align="center" valign="top"> GU237693
</td><td align="left" valign="top"><italic>Chrysanthemum parthenii</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"> PD 85/259
</td><td align="left" valign="top"><italic>Stagonosporopsis ligulicola</italic> var. <italic>inoxydabilis</italic></td><td align="center" valign="top"> GU238189
</td><td align="center" valign="top"> GU237920
</td><td align="center" valign="top"> GU237694
</td><td align="left" valign="top"><italic>Matricaria</italic> sp.
</td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=500.63&link_type=cbs">CBS 500.63</ext-link>; MUCL
8090
</td><td align="left" valign="top"><italic>Stagonosporopsis ligulicola</italic> var. <italic>ligulicola</italic> B
</td><td align="center" valign="top"> GU238190
</td><td align="center" valign="top"> GU237872
</td><td align="center" valign="top"> GU237695
</td><td align="left" valign="top"><italic>Chrysanthemum indicum</italic></td><td align="left" valign="top"> Germany
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=137.96&link_type=cbs">CBS 137.96</ext-link>; PD
84/75
</td><td align="left" valign="top"><italic>Stagonosporopsis ligulicola</italic> var. <italic>ligulicola</italic> B
</td><td align="center" valign="top"> GU238191
</td><td align="center" valign="top"> GU237783
</td><td align="center" valign="top"> GU237696
</td><td align="left" valign="top"><italic>Chrysanthemum indicum</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=562.81&link_type=cbs">CBS 562.81</ext-link>; PDDCC
6884
</td><td align="left" valign="top"><italic>Stagonosporopsis loticola</italic> T
</td><td align="center" valign="top"> GU238192
</td><td align="center" valign="top"> GU237890
</td><td align="center" valign="top"> GU237697
</td><td align="left" valign="top"><italic>Lotus pedunculatus</italic></td><td align="left" valign="top"> New Zealand
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=628.97&link_type=cbs">CBS 628.97</ext-link>; PD
79/72; PDDCC 3870
</td><td align="left" valign="top"><italic>Stagonosporopsis loticola</italic></td><td align="center" valign="top"> GU238193
</td><td align="center" valign="top"> GU237896
</td><td align="center" valign="top"> GU237698
</td><td align="left" valign="top"><italic>Lotus tenuis</italic></td><td align="left" valign="top"> New Zealand
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=101494&link_type=cbs">CBS 101494</ext-link>; PD
98/5247
</td><td align="left" valign="top"><italic>Stagonosporopsis lupini</italic> B
</td><td align="center" valign="top"> GU238194
</td><td align="center" valign="top"> GU237724
</td><td align="center" valign="top"> GU237699
</td><td align="left" valign="top"><italic>Lupinus albus</italic></td><td align="left" valign="top"> U.K.
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=375.84&link_type=cbs">CBS 375.84</ext-link>; PD
80/1250
</td><td align="left" valign="top"><italic>Stagonosporopsis lupini</italic></td><td align="center" valign="top"> GU238195
</td><td align="center" valign="top"> GU237844
</td><td align="center" valign="top"> GU237700
</td><td align="left" valign="top"><italic>Lupinus mutabilis</italic></td><td align="left" valign="top"> Peru
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=634.92&link_type=cbs">CBS 634.92</ext-link>; IMI
193307
</td><td align="left" valign="top"><italic>Stagonosporopsis oculo-hominis</italic> T
</td><td align="center" valign="top"> GU238196
</td><td align="center" valign="top"> GU237901
</td><td align="center" valign="top"> GU237701
</td><td align="left" valign="top"> Human
</td><td align="left" valign="top"> U.S.A.
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=109180&link_type=cbs">CBS 109180</ext-link>; PD
79/175
</td><td align="left" valign="top"><italic>Stagonosporopsis rudbeckiae</italic> B
</td><td align="center" valign="top"> GU238197
</td><td align="center" valign="top"> GU237745
</td><td align="center" valign="top"> GU237702
</td><td align="left" valign="top"><italic>Rudbeckia bicolor</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=379.91&link_type=cbs">CBS 379.91</ext-link>; PD
77/675
</td><td align="left" valign="top"><italic>Stagonosporopsis trachelii</italic> B
</td><td align="center" valign="top"> GU238173
</td><td align="center" valign="top"> GU237850
</td><td align="center" valign="top"> GU237678
</td><td align="left" valign="top"><italic>Campanula isophylla</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=384.68&link_type=cbs">CBS 384.68</ext-link></td><td align="left" valign="top"><italic>Stagonosporopsis trachelii</italic> B
</td><td align="center" valign="top"> GU238174
</td><td align="center" valign="top"> GU237856
</td><td align="center" valign="top"> GU237679
</td><td align="left" valign="top"><italic>Campanula isophylla</italic></td><td align="left" valign="top"> Sweden
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=273.92&link_type=cbs">CBS 273.92</ext-link>; PD
76/1019
</td><td align="left" valign="top"><italic>Stagonosporopsis valerianellae</italic></td><td align="center" valign="top"> GU238200
</td><td align="center" valign="top"> GU237819
</td><td align="center" valign="top"> GU237705
</td><td align="left" valign="top"><italic>Valerianella locusta</italic></td><td align="left" valign="top"> Netherlands
</td></tr><tr><td align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=329.67&link_type=cbs">CBS 329.67</ext-link>; PD
66/302
</td><td align="left" valign="top"><italic>Stagonosporopsis valerianellae</italic> B
</td><td align="center" valign="top">GU238201
</td><td align="center" valign="top">GU237832
</td><td align="center" valign="top">GU237706
</td><td align="left" valign="top"><italic>Valerianella locusta</italic> var. <italic>oleracea</italic></td><td align="left" valign="top">Netherlands
</td></tr></tbody></table><table-wrap-foot><fn id="tblfn4"><label>1</label><p>ATCC: American Type Culture Collection, Virginia, U.S.A.; CBS:
Centraalbureau voor Schimmelcultures, Utrecht, The Netherlands; CECT:
Colección Española de Cultivos Tipo, Valencia University, Spain;
DAOM: Canadian Collection of Fungal Cultures, Ottawa, Canada; DSM: Deutsche
Sammlung von Mikroorganismen und Zellkulturen GmbH, Braunschweig, Germany;
HACC: Research Laboratory, Hindustan Antibiotics Ltd., Pimpri Poona, India;
IMI: International Mycological Institute, CABI-Bioscience, Egham, Bakeham
Lane, U.K.; LEV: Plant Health and Diagnostic Station, Auckland, New Zealand;
MUCL: Mycotheque de l'Universite catholique de Louvain, Louvain-la-Neuve,
Belgium; PD: Plant Protection Service, Wageningen, the Netherlands; PDDCC:
Plant Diseases Division Culture Collection, Auckland, New Zealand; PREM:
National Collection of Fungi: Culture Collection, Pretoria, South Africa; VKM:
All-Russian Collection of Microorganisms, Pushchino, Russia; VPRI: Victorian
Plant Disease Herbarium, Victoria, Australia.</p></fn><fn id="tblfn5"><label>2</label><p>T: Ex-type strain; B: Reference strain according to Boerema <italic>et al.</italic>(<xref ref-type="bibr" rid="ref55">2004</xref>).</p></fn></table-wrap-foot></table-wrap></p><p>Obtained consensus sequences were assembled and aligned using the same
BioNumerics software and adjusted manually where necessary. As SSU was highly
conserved in deeper node phylogenies, revealing almost no phylogenetic
informative nuclear polymorphisms, and as ITS and TUB proved to be unalignable
due to a high level of polymorphism if all taxa studied would be taken into
account, it was decided to conduct two separate analyses. The first analysis
comprised SSU and LSU loci, and was applied to 76 taxa of which most species
included belonged to genera that were often confused with <italic>Phoma</italic>(<xref ref-type="bibr" rid="ref219">Sutton 1980</xref>,
<xref ref-type="bibr" rid="ref100">De Gruyter <italic>et al.</italic>2009</xref>). A second set of analyses was conducted on 274 taxa, and
focussed on the species that had proven to be related to the
<italic>Didymellaceae</italic> from preliminary studies.</p><p>Each of the phylogenetic analyses consisted of two methods: Bayesian
Interference (BI) and Maximum Likelihood (ML). For BI analysis, the nucleotide
substitution models were determined for each locus separately with MrModeltest
v. 2.2 (<xref ref-type="bibr" rid="ref153">Nylander 2004</xref>).
According to this software, the General Time Reversible substitution was
determined to be the best model for SSU, TUB and LSU in both data sets, with
inverse gamma rates and dirichlet base frequencies (GTR + I + G). For the ITS
dataset, the software suggested the Symmetrical Model as the best model for
substitution of nucleotides. Also in this locus, the inverse gamma rates and
dirichlet base frequencies were used (SYM + I + G). The actual Bayesian
calculations were performed in MrBayes v. 3.1.2
(<xref ref-type="bibr" rid="ref113">Huelsenbeck & Ronquist
2001</xref>). One tree was saved per 100 generations, and the run was
automatically ended when the standard deviation of split frequencies was below
0.01. The temperature value of the Bayesian run was set at 0.2. To avoid
suboptimal trees being taking into account for the consensus tree, a burn-in
of 25 % of the saved trees was used. The resulting “50 % majority rule
consensus” trees were visualised with TreeView v. 1.6.6
(<xref ref-type="bibr" rid="ref157">Page 1996</xref>).</p><p>A second measure of branch support was obtained by conducting a ML analysis
using RAxML software (<xref ref-type="bibr" rid="ref210">Stamatakis <italic>et
al.</italic> 2005</xref>) through the CIPRES Website
(<ext-link ext-link-type="uri" xlink:href="www.phylo.org">www.phylo.org</ext-link>).
The same partitions were used as in the BI analyses, but because RAxML
implements only the GTR substitution model, the symmetrical model for the ITS
partition was waived. The robustness of trees in the ML analyses was evaluated
by bootstrapping the datasets. The number of bootstrap replicates was
automatically determined by the RAxML software
(<xref ref-type="bibr" rid="ref211">Stamatakis <italic>et al.</italic>2008</xref>). The obtained trees in both analyses are lodged with
TreeBASE
(<ext-link ext-link-type="uri" xlink:href="www.treebase.org">www.treebase.org</ext-link>).</p></sec><sec><title>Morphology</title><p>Morphological studies of the strains were performed on OA, malt extract
agar (MEA) and cherry decoction agar (CHA)
(<xref ref-type="bibr" rid="ref76">Crous <italic>et al</italic>. 2009c</xref>).
The cultures were incubated according to the methodologies described by
Boerema <italic>et al</italic>.
(<xref ref-type="bibr" rid="ref55">2004</xref>). Eight days after
inoculation, the colony growth was measured. At the 15<sup>th</sup> day after
incubation, the colony colours were rated using the colour charts of Rayner
(<xref ref-type="bibr" rid="ref177">1970</xref>). Micromorphological
features were studied after maturation of the pycnidia. Therefore, fungal
structures were mounted in tap water using a scalpel blade and examined under
a stereo light microscope. Perennial structures that were formed in the agar
medium, such as chlamydospores, were cut out from the medium, and mounted in
lactic acid. Remaining agar was removed from these samples by gently heating
the glass slides. The sizes of the various structures were determined by
averaging the measurements of 30 samples of each structure, except for
conidiogenous cells and pycnidial wall characters, of which the size ranges
were estimated based on 5–10 samples. Fifth and 95<sup>th</sup>percentiles were determined for all measurements and are provided in
parentheses. By application of a droplet of 1N NaOH, the production of
metabolite E+ was determined (<xref ref-type="bibr" rid="ref83">Dorenbosch
1970</xref>, <xref ref-type="bibr" rid="ref152">Noordeloos <italic>et
al</italic>. 1993</xref>). The structure of the pycnidial wall and shape of
conidiogenous cells were studied using microtome sections of 6 μm
thickness, prepared with a Leica CM3050 freezing microtome and mounted in
lactic acid. Taxonomic recombinations and novel species and descriptions were
deposited in MycoBank.</p></sec></sec><sec><title>RESULTS</title><sec><title>Systematics of the genus <italic>Phoma</italic></title><sec><title>DNA phylogenetical analysis</title><p>Due to alignment difficulties multiple datasets, consisting of different
sets of loci, were utilised. For a generic overview, LSU and SSU were included
in the first alignment, which consisted of 76 taxa. A list of species names
and numbers, original substrates, geographical origins and GenBank accession
numbers of the strains used in this study is provided in
<xref ref-type="table" rid="tbl2">Table 2</xref>. The aligned sequence
matrix had a total length of 2 210 characters including alignment gaps (LSU: 1
258 and SSU: 952 bp). Of those characters, 1 809 (LSU: 994 and SSU: 815) were
constant and 401 were variable (LSU: 264 and SSU: 137). The Bayesian analysis
run was aborted after 10 000 000 generations as a point of stationarity was
reached in the average standard deviation of split frequencies, at a value of
0.0288. The applied “burn-in” percentage of 25 % was well after
stationarity in the probability of the trees was reached. The tree topologies
and support values of the ML analysis, differed only slightly from the trees
obtained from the Bayesian analyses, supporting the probability of the tree.
The tree is rooted to <italic>Pseudorobillarda phragmitis</italic>(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=398.61&link_type=cbs">CBS 398.61</ext-link>).</p><p>Based on the LSU-SSU phylogenetic study performed here for the various
anamorph and teleomorph species in the <italic>Phoma</italic> complex, eight clades
were revealed (<xref ref-type="fig" rid="fig1">Fig. 1</xref>),
including one which only comprises the outgroup specimen. The various clades
will be treated below, but for additional synonymy on the <italic>Phoma</italic>species we refer to Boerema <italic>et al.</italic>(<xref ref-type="bibr" rid="ref55">2004</xref>). The findings in these
clades are largely in congruence with the observations of De Gruyter <italic>et
al.</italic> (<xref ref-type="bibr" rid="ref100">2009</xref>).</p><p>Species that were ascribed to the <italic>Phoma</italic> section <italic>Phoma</italic> by
Boerema <italic>et al.</italic> (<xref ref-type="bibr" rid="ref55">2004</xref>)
appear to be genetically highly heterogeneous, as these species are recovered
in almost every clade. Species that were ascribed to <italic>Phoma</italic> section
<italic>Heterospora</italic> appear to be linked to at least three distinct clades.
Also polymorphism is observed for sections <italic>Paraphoma, Peyronellaea</italic>and <italic>Sclerophomella</italic>, as well as for <italic>Coniothyrium</italic> and
<italic>Ascochyta</italic>. The type species of this latter genus, <italic>A. pisi</italic>,
is not included in the present tree, but is genetically similar to the
<italic>Didymellaceae</italic>.</p><p><fig position="float" id="fig1"><label>Fig. 1.</label><caption><p>(p. 15) Fifty percent majority rule consensus tree from a BI analysis of
Large and Small subunit sequences of <italic>Phoma</italic> and related genera (n =
76). At the nodes the BI Posterior Probabilities are presented above the
branch, and bootstrap percentages of the ML analysis are given below the
branch. Branches that were less than 50 % supported in the ML analyses are
indicated with a hyphen. The bar indicates the number of substitutions per
site. The tree is rooted with <italic>Pseudorobillarda phragmitis</italic>(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=398.61&link_type=cbs">CBS 398.61</ext-link>).</p></caption><graphic xlink:href="1fig1"></graphic></fig></p></sec><sec><title>Treatment of the clades</title><sec><title>Clade 1, Outgroup:</title><p><italic>Pseudorobillarda phragmitis</italic> was selected as outgroup on the basis
of the studies conducted by De Gruyter <italic>et al</italic>.
(<xref ref-type="bibr" rid="ref100">2009</xref>). This species,
although being recognised as a coelomycete, is not only phylogenetically, but
also morphologically distinct from <italic>Phoma</italic>, although Sutton
(<xref ref-type="bibr" rid="ref219">1980</xref>) classified it in the
Phialopycnidiineae.</p></sec><sec><title>Clade 2, Sporormiaceae:</title><p>In the basal lineages, <italic>Sporormiella minima</italic>(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=524.50&link_type=cbs">CBS 524.50</ext-link>) was
recovered, representing the <italic>Sporormiaceae</italic>, which was recently
recircumscribed (<xref ref-type="bibr" rid="ref22">Barr 2000</xref>).
In the same clade, two species were recovered that are described in
<italic>Phoma</italic> section <italic>Phoma</italic>: <italic>Ph. capitulum</italic> and <italic>Ph.
minutispora</italic>. Both species are distinguishable from other species in this
Boeremaean section by the production of relatively small subglobose conidia
(measuring <italic>ca</italic>. 2–5 × 1.5–3 μm) with a few, large
guttules. Within the <italic>Sporormiaceae</italic>, teleomorphs species have been
reported with phoma-like anamorphs, such as <italic>Westerdykella dispersa</italic>(<xref ref-type="bibr" rid="ref14">Von Arx 1981</xref>). Two
<italic>Sporormiaceae</italic>-associated genera, <italic>Sporormia</italic> and
<italic>Preussia</italic>, have been mentioned as possible teleomorph for <italic>Ph.
deserticola</italic> (<xref ref-type="bibr" rid="ref16">Von Arx & Storm
1967</xref>), a species that was regarded as miscellaneous by Boerema
<italic>et al.</italic> (<xref ref-type="bibr" rid="ref55">2004</xref>). Also
these anamorphs produce minute (sub-) globose conidia
(<xref ref-type="bibr" rid="ref14">Von Arx 1981</xref>,
<xref ref-type="bibr" rid="ref55">Boerema <italic>et al.</italic> 2004</xref>).
Although the <italic>Sporormiaceae</italic> belongs to the <italic>Pleosporales</italic> (Barr
<xref ref-type="bibr" rid="ref22">2000</xref>,
<xref ref-type="bibr" rid="ref23">2002</xref>,
<xref ref-type="bibr" rid="ref201">Shearer <italic>et al</italic>. 2009</xref>,
<xref ref-type="bibr" rid="ref213">Suetrong <italic>et al.</italic>2009</xref>), it forms a rather basal clade to most of the other
<italic>Phoma</italic> species, and a taxonomic revision of <italic>Ph. capitulum</italic> and
<italic>Ph. minutispora</italic> should therefore be considered.</p></sec><sec><title>Clade 3, Melanommataceae:</title><p>One species that belongs to the <italic>Melanommataceae</italic> was included in
the phylogenetical reconstruction of the phomoid <italic>Pleosporales</italic>. This
species, <italic>Aposphaeria populina</italic>(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=543.70&link_type=cbs">CBS 543.70</ext-link>), is
recovered in the basal lineages of the reconstructed tree
(<xref ref-type="bibr" rid="ref141">Mugambi & Huhndorf 2009</xref>,
<xref ref-type="bibr" rid="ref213">Suetrong <italic>et al.</italic>2009</xref>, <xref ref-type="bibr" rid="ref222">Tanaka <italic>et al</italic>.
2009</xref>). The close association of this family with the
<italic>Sporormiaceae</italic> and their phylogenetic placement in the basal lineages
of the <italic>Pleosporales</italic> is in congruence with results obtained in earlier
studies (<xref ref-type="bibr" rid="ref130">Kruys <italic>et al.</italic>2006</xref>, <xref ref-type="bibr" rid="ref100">De Gruyter <italic>et
al.</italic> 2009</xref>). Although some earlier workers regularly mistook
several <italic>Phoma</italic> species for members of the genus <italic>Aposphaeria</italic>(<italic>e.g.</italic> <xref ref-type="bibr" rid="ref188">Saccardo
1884</xref>), none of the <italic>Phoma</italic> species included in this study
were clustering with the <italic>Melanommataceae</italic>.</p></sec><sec><title>Clade 4:</title><p>This clade comprises a range of species that almost all belong to different
genera. <italic>Phoma lini</italic> and <italic>Ph. flavescens</italic> are the two
<italic>Phoma</italic> representatives found in this clade, although they are not
sister species. Based on morphological data, both species were accommodated in
<italic>Phoma</italic> section <italic>Phoma</italic> (<xref ref-type="bibr" rid="ref97">De
Gruyter <italic>et al</italic>. 1993</xref>). Both species produce a yellow
diffusible pigment <italic>in vitro</italic>, although a positive reaction to NaOH is
only observed in <italic>Ph. lini</italic>. Both <italic>Ph. flavescens</italic> and <italic>Ph.
lini</italic> are closely related to <italic>Paraconiothyrium minitans</italic> (≡
<italic>Coniothyrium minitans</italic>; <xref ref-type="bibr" rid="ref228">Verkley
<italic>et al.</italic> 2004</xref>). With this formal recombination into
<italic>Paraconiothyrium</italic>, it was aimed to differentiate <italic>Par.
minitans</italic>, which produces complex, thick-walled pycnidia from other
<italic>Coniothyrium</italic> species that normally produce more phomoid pycnidia
(<xref ref-type="bibr" rid="ref228">Verkley <italic>et al.</italic>2004</xref>). The close relationship between <italic>Par. minitans</italic> with
<italic>C. fuckelii</italic> that is found here is in congruence with the observations
of Damm <italic>et al.</italic>(<xref ref-type="bibr" rid="ref78">2008</xref>), although the
teleomorph name, <italic>Leptosphaeria coniothyrium</italic>, would suggest a
association with the <italic>Leptosphaeriaceae</italic> (clade 8).</p><p>The likeliness of the findings of <italic>Pyrenochaeta romeroi</italic>(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=252.60&link_type=cbs">CBS 252.60</ext-link>),
<italic>Asteromella tiliae</italic>(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=265.94&link_type=cbs">CBS 265.94</ext-link>) and
<italic>Neottiosporina paspali</italic>(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=331.37&link_type=cbs">CBS 331.37</ext-link>) in
this clade was already discussed by De Gruyter <italic>et al.</italic>(<xref ref-type="bibr" rid="ref100">2009</xref>).</p></sec><sec><title>Clade 5, Cucurbitariaceae:</title><p>Clade 5 comprises mainly taxa with setose pycnidia, including several
representative species of the genus <italic>Pyrenochaeta.</italic> In addition, a
<italic>Coniothyrium</italic> sp., <italic>Phialophorophoma litoralis</italic> and
<italic>Pleurophoma cava</italic> grouped in this clade, as well as two <italic>Phoma</italic>species, <italic>Ph. pratorum</italic> (section <italic>Phoma</italic>) <italic>and Ph.
terricola</italic>, (section <italic>Paraphoma</italic>). Another representative of the
section <italic>Paraphoma</italic> that is included in this study is <italic>Ph.
radicina</italic>, which is however found in clade 6. The taxonomy of setose
species that are currently classified in <italic>Phoma</italic> section <italic>Paraphoma,
Pyrenochaeta</italic> and <italic>Pleurophoma</italic> is revised
(<xref ref-type="bibr" rid="ref101">De Gruyter <italic>et al.</italic>2010</xref>). Also in several non-<italic>Paraphoma</italic> species in the genus
<italic>Phoma</italic> setose or semi-pilose pycnidia do occur incidentally
(<xref ref-type="bibr" rid="ref55">Boerema <italic>et al.</italic>2004</xref>). However, thus far, no setae-forming <italic>Ph. pratorum</italic>strains have been recorded. The finding of this species in the present clade
is thus highly remarkable.</p><p>The <italic>Coniothyrium</italic> strain in this clade
(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=122787&link_type=cbs">CBS 122787</ext-link>) was
previously identified as <italic>C. cerealis</italic>, and is found to be closely
related to <italic>Pyrenochaeta acicola</italic> (BPP = 0.99, RBS = 100 %). As was
illustrated in a previous study of Muthumeenakshi <italic>et al.</italic>(<xref ref-type="bibr" rid="ref145">2001</xref>) <italic>C. cerealis</italic>is quite distantly related to other <italic>Coniothyrium</italic> species. However,
based on comparison with sequence data available in GenBank, it is unlikely
that its previous identification was correct. This finding further illustrates
the polyphyly of the genus <italic>Coniothyrium</italic>, which further has been
retrieved in clades 4, 6 (<italic>Phaeosphaeriaceae</italic>), 7
(<italic>Leptosphaeriaceae</italic> and <italic>Pleosporaceae</italic>) and 8
(<italic>Didymellaceae</italic>). As mentioned before, some species of this genus have
been associated with the teleomorph genus <italic>Leptosphaeria</italic>, and are thus
expected to cluster with the <italic>Leptosphaeriaceae</italic> (clade 7). None of the
species recovered in clade 5 has been associated with a teleomorph.</p></sec><sec><title>Clade 6, Phaeosphaeriaceae:</title><p>The species that are found in the well-supported clade 6 (BPP = 1.00; RBS =
83 %), belong to the morphologically heterogeneous group of the
<italic>Phaeosphaeriaceae</italic> (<xref ref-type="bibr" rid="ref28">Boehm <italic>et
al</italic>. 2009</xref>, <xref ref-type="bibr" rid="ref242">Zhang <italic>et
al.</italic> 2009</xref>). Most findings in this clade have already been
discussed in the previous paper of De Gruyter <italic>et al.</italic>(<xref ref-type="bibr" rid="ref100">2009</xref>). In addition to that
study, six <italic>Phoma</italic> species are retrieved in this clade. <italic>Phoma
radicina</italic>, type of <italic>Phoma</italic> section <italic>Paraphoma</italic>, is found in
close association with <italic>Ph. chrysanthemicola</italic> (BPP = 1.00; RBS = 90 %).
The association between <italic>Ph. radicina</italic> and the
<italic>Phaeosphaeriaceae</italic> is further discussed by De Gruyter <italic>et al.</italic>(<xref ref-type="bibr" rid="ref101">2010</xref>). Its close association
with <italic>Ph. chrysanthemicola</italic> has been observed before by Aveskamp <italic>et
al.</italic> (2008a), but the link with the <italic>Phaeosphaeriaceae</italic> has not
been established. Strains of <italic>Ph. chrysanthemicola</italic> exhibit some
semi-setose pycnidia that are, however, often fully covered by mycelial hairs
(<xref ref-type="bibr" rid="ref33">Boerema 1993</xref>). This is a
feature that is in common with <italic>Ph. radicina</italic>, which has, as type
species of the section <italic>Paraphoma</italic>, clearly visible setae. In contrast,
the main characteristic of <italic>Ph. chrysanthemicola</italic>, the presence of
pseudosclerotioid masses, has never been observed in the latter species.
However, also not all strains of <italic>Ph. chrysanthemicola</italic> exhibit this
character (<xref ref-type="bibr" rid="ref83">Dorenbosch
1970</xref>).</p><p><italic>Phoma fimeti</italic> forms a subclade with <italic>Ph. samarorum</italic> and a
strain that was previously identified as <italic>Stenocarpella macrospora</italic>(BPP = 0.98; RBS = 67 %), but that is probably misidentified
(<xref ref-type="bibr" rid="ref100">De Gruyter <italic>et al.</italic>2009</xref>). Especially the finding of <italic>Ph. samarorum</italic> is
noteworthy, as it is found rather distinct from two clusters of other species
belonging to the section <italic>Heterospora</italic>, which are retrieved among the
<italic>Leptosphaeriaceae</italic> and <italic>Didymellaceae</italic> (clades 7 and 8). In
contrast to these other <italic>Heterospora</italic> species, the large conidia of
<italic>Ph. samarorum</italic> that can be observed <italic>in planta</italic> are clearly
distinct by the subulate top cells, and measures up to 17 × 3.5 μm
(<xref ref-type="bibr" rid="ref54">Boerema <italic>et al.</italic>1997</xref>). The strain identified as <italic>Stenocarpella macrospora</italic>is now sterile and therefore not studied morphologically. This species is
known to produce similar-shaped, septate conidia, which are however pigmented
and considerably larger, 44–82 × 7.5–11.5 μm
(<xref ref-type="bibr" rid="ref219">Sutton 1980</xref>). The close
association with <italic>Ph. fimeti</italic> is therefore remarkable as this species
is known to produce only minute, aseptate conidia, measuring
(2–)2.5–4(–5) × (1.5–)2–2.5(–3)
μm (<xref ref-type="bibr" rid="ref96">De Gruyter & Noordeloos
1992</xref>).</p><p>The remaining two <italic>Phoma</italic> species in this clade, <italic>Ph.
haematocycla</italic> and <italic>Ph. opuntiae</italic>, also produce such minute conidia.
<italic>Phoma haematocyla</italic>, a flax-associated species from New Zealand, is
retrieved in a subclade that also accommodates <italic>Chaetasphaeronema
hispidulum</italic> (BPP = 1.00; RBS = 100 %).</p><p>All <italic>Phoma</italic> species found here are morphologically rather distinct,
hence their placement in four different <italic>Phoma</italic> sections
(<xref ref-type="bibr" rid="ref55">Boerema <italic>et al.</italic>2004</xref>). None of the <italic>Phoma</italic> species accommodated in this
clade is associated with a teleomorph. The main teleomorph associated with the
<italic>Phaeosphaeriaceae</italic> is <italic>Phaeosphaeria,</italic> although also
incidentally a <italic>Leptosphaeria</italic> species is associated with this family
(<xref ref-type="bibr" rid="ref62">Câmara <italic>et al.</italic>2002</xref>). An anamorph genus that is often confused with
<italic>Phoma</italic> is <italic>Microsphaeropsis</italic>(<xref ref-type="bibr" rid="ref34">Boerema 1997</xref>), which is
linked to <italic>Phaeosphaeria</italic>(<xref ref-type="bibr" rid="ref62">Câmara <italic>et al.</italic>2002</xref>). Both anamorph genera differ in conidial pigmentation, which
is commonly only present in mature conidia of <italic>Microsphaeropsis</italic>.
Younger conidia are, however, often colourless. It may be that the
<italic>Phoma</italic> species in this clade actually belong to what is now known as
<italic>Microsphaeropsis</italic>, but have lost the pigmentation character during
evolution.</p></sec><sec><title>Clade 7, Leptosphaeriaceae and Pleosporaceae:</title><p>Clade 7 is a large clade comprising many <italic>Phoma</italic> species from
various Boeremaean sections. Three reference species encountered here have
been associated with the <italic>Leptosphaeriaceae</italic> before, these include
<italic>Leptosphaeria maculans, L. biglobosa</italic> and <italic>Coniothyrium
palmarum</italic> (<xref ref-type="bibr" rid="ref178">Reddy <italic>et al.</italic>1998</xref>, <xref ref-type="bibr" rid="ref228">Verkley <italic>et al.</italic>2004</xref>, <xref ref-type="bibr" rid="ref100">De Gruyter <italic>et
al.</italic> 2009</xref>), or with the <italic>Pleosporaceae</italic>, such as
<italic>Pleospora herbarum, Ascochyta caulina</italic> and <italic>Ph. betae</italic>(<xref ref-type="bibr" rid="ref82">Dong <italic>et al.</italic> 1998</xref>,
<xref ref-type="bibr" rid="ref126">Kodsueb <italic>et al.</italic> 2006</xref>,
<xref ref-type="bibr" rid="ref116">Inderbitzin <italic>et al.</italic>2009</xref>, <xref ref-type="bibr" rid="ref100">De Gruyter <italic>et
al.</italic> 2009</xref>).</p><p>The two <italic>Leptosphaeria</italic> species in this study that were associated
with a <italic>Phoma</italic> anamorph cluster together in the present clade: <italic>L.
maculans</italic> (anam <italic>Ph. lingam</italic>) and <italic>L. biglobosa</italic>, which
produces an unnamed, phomoid anamorph that is highly similar to <italic>Ph.
lingam</italic> (<xref ref-type="bibr" rid="ref202">Shoemaker & Brun
2001</xref>). Both species are serious pathogens of <italic>Brassicaceae</italic>(<xref ref-type="bibr" rid="ref89">Fitt <italic>et al.</italic> 2006</xref>).
<italic>Leptosphaeria biglobosa</italic> was found to be closely related to <italic>Ph.
lingam</italic> in previous studies (Mendes-Perreira <italic>et al.</italic> 2003) and was
for a long time recognised as a weakly pathogenic variety of the latter
species (<xref ref-type="bibr" rid="ref118">Johnson & Lewis
1990</xref>, <xref ref-type="bibr" rid="ref191">Schäfer &
Wöstemeyer 1992</xref>, <xref ref-type="bibr" rid="ref138">Morales
<italic>et al</italic>. 1993</xref>, <xref ref-type="bibr" rid="ref167">Pongam
<italic>et al</italic>. 1999</xref>,
<xref ref-type="bibr" rid="ref235">Williams & Fitt 1999</xref>,
<xref ref-type="bibr" rid="ref173">Purwantara <italic>et al</italic>.
2000</xref>, <xref ref-type="bibr" rid="ref202">Shoemaker & Brun
2001</xref>, <xref ref-type="bibr" rid="ref230">Voigt <italic>et al</italic>.
2001</xref>).</p><p>The phylogenic relation of <italic>Phoma</italic> species currently classified in
sections <italic>Pleonodomus</italic> and <italic>Pilosa</italic> is currently investigated
(De Gruyter <italic>et al</italic>. in prep.). However, the present results reveal
that a number of species from other <italic>Phoma</italic> sections fits in the
<italic>Leptosphaeriaceae</italic> and <italic>Pleosporaceae</italic>. These include <italic>Ph.
apiicola, Ph. fallens, Ph. flavigena, Ph. glaucispora, Ph. multipora, Ph.
valerianeae</italic> and <italic>Ph. vasinfecta</italic>. In contrast to the species that
are accommodated in sections <italic>Pilosa</italic> and <italic>Plenodomus</italic>, pilose
or scleroplectenchymatous pycnidia have never been recorded in these seven
species; hence the placements in section <italic>Phoma</italic>.</p><p><italic>Phoma multipora</italic> was ascribed to section <italic>Phoma</italic>. However,
the original morphological description mentions the presence of elongated
conidiophores (<xref ref-type="bibr" rid="ref160">Pawar <italic>et al.</italic>1967</xref>), which indicates that this species does not belong to the
genus <italic>Phoma</italic> according to the present-day concept.</p><p>In addition, some representatives of other sections are found in clade 7,
such as <italic>Ph. incompta</italic> (section <italic>Sclerophomella</italic>) and <italic>Ph.
violicola</italic>, which is associated with the section <italic>Peyronellaea</italic>.
Based on previous studies in the section <italic>Peyronellaea</italic> however, also
<italic>Ph. chrysanthemicola</italic> and <italic>Ph. schachtii</italic> may be expected to
cluster with the species in this clade
(<xref ref-type="bibr" rid="ref19">Aveskamp <italic>et al</italic>.
2009a</xref>). Remarkably, also two representatives of the section
<italic>Heterospora</italic> are found in this clade. <italic>Phoma heteromorphospora</italic>is the assigned type species of this section
(<xref ref-type="bibr" rid="ref54">Boerema <italic>et al.</italic>1997</xref>), whereas <italic>Ph. dimorphospora</italic> is morphologically
closely allied, in congruence with the molecular results obtained here. Both
species have a slow growth-rate and occur on <italic>Chenopodium</italic> spp., but
can be distinguished by the absence of the conidial dimorphism in <italic>Ph.
dimorphospora in vitro</italic>. Moreover, the latter species is commonly found in
North and South America, whilst <italic>Ph. heteromorphospora</italic> occurs mainly
in Europe (<xref ref-type="bibr" rid="ref55">Boerema <italic>et al</italic>.
2004</xref>).</p><p>With the exception of <italic>Ph. samarorum</italic> (clade 6 –
<italic>Phaeosphaeriaceae</italic>), the other species of the section
<italic>Heterospora</italic> are found in clade 8, which represents the
<italic>Didymellaceae</italic>. The major difference between the <italic>Heterospora</italic>species in the present clade in contrast to those in the
<italic>Didymellaceae</italic> is the size of the septate conidia, which are up to 9
× larger <italic>in vivo</italic> than the regular conidia in <italic>Ph.
heteromorphospora</italic> and <italic>Ph. dimorphospora</italic>, whereas, in the
<italic>Didymellaceae</italic> clade, the septate conidia are only 1.5–4.5
× larger.</p><p>Also, <italic>Coniothyrium palmarum</italic>, which represents the type of its
genus, clusters in this clade. Just as in <italic>Phoma</italic>, the species in
<italic>Coniothyrium</italic> have only a limited number of morphological features
that can aid in taxonomy. This has led to an unwanted situation in which
species morphologically placed in this genus have been shown in phylogenetic
examination to be dispersed among multiple families
(<xref ref-type="bibr" rid="ref228">Verkley <italic>et al</italic>.
2004</xref>). Although, based on type species, an anamorph-teleomorph
link has been established between <italic>Coniothyrium</italic> and
<italic>Leptosphaeria</italic> (<xref ref-type="bibr" rid="ref71">Crous
1998</xref>), many heterogeneous species are <italic>Coniothyrium</italic>-like,
and belong phylogenetically to different families or even classes
(<xref ref-type="bibr" rid="ref70">Cortinas <italic>et al</italic>.
2006</xref>). In this study we found “<italic>Coniothyrium</italic>”
species accommodated in at least three different clades
(<xref ref-type="fig" rid="fig1">Fig. 1</xref>). <italic>Coniothyrium
clematidis-rectae</italic> is phylogenetically linked to the
<italic>Didymellaceae</italic> (<xref ref-type="fig" rid="fig2">Fig. 2</xref>– see below). <italic>Phoma</italic> and <italic>Coniothyrium</italic> are considered to
be highly similar and are only distinguished on basis of the pigmentation of
the conidia and the structure of the pycnidial wall
(<xref ref-type="bibr" rid="ref55">Boerema <italic>et al</italic>.
2004</xref>).</p><p>This clade also accommodates <italic>Pleospora betae</italic>, a notorious leaf and
seed pathogen of beet (<italic>Beta vulgaris</italic>,
<xref ref-type="bibr" rid="ref61">Bugbee & Cole 1981</xref>), and
<italic>Pl. herbarum</italic>, which is the type species of the genus
<italic>Pleospora</italic>. The genetic distance between the two species was already
observed in a study utilising SSU nrDNA sequences
(<xref ref-type="bibr" rid="ref82">Dong <italic>et al</italic>. 1998</xref>).
Also three <italic>Phoma</italic> species that are found in close association with
these “true” <italic>Pleosporaceae</italic> and that are found basal to
this clade, <italic>Ph. fallens, Ph. flavigena</italic> and <italic>Ph. glaucispora</italic>have glabrous pycnidia and, like <italic>Ph. betae</italic>, aseptate conidia, hence
their link to <italic>Phoma</italic> section <italic>Phoma</italic>. Absence of an ostiole is
only recorded in <italic>Ph. glaucispora</italic>(<xref ref-type="bibr" rid="ref98">De Gruyter <italic>et al</italic>.
1998</xref>).</p><p><fig position="float" id="fig2"><label>Fig. 2.</label><caption><p>Fifty percent majority rule consensus tree from a BI analysis of LSU, ITS
and TUB sequences of <italic>Didymellaceae</italic> (n = 274). At the nodes the BI
Posterior Probabilities are presented above the branch, and bootstrap
percentages of the analysis are given below the branch. Branches that were
less than 50 % supported in the ML analyses are indicated with a hyphen. The
bar indicates the number of substitutions per site. The tree is rooted with
<italic>Ascochyta hordei</italic> var. <italic>hordei</italic>(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=544.74&link_type=cbs">CBS 544.74</ext-link>) and
<italic>Phoma paspali</italic> (<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=560.81&link_type=cbs">CBS
560.81</ext-link> & <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=561.81&link_type=cbs">CBS
561.81</ext-link>).></p></caption><graphic xlink:href="1fig2A"></graphic><graphic xlink:href="1fig2B"></graphic><graphic xlink:href="1fig2C"></graphic><graphic xlink:href="1fig2D"></graphic></fig></p><p><italic>Pleospora</italic> is linked to the anamorph genus <italic>Stemphylium</italic>(<xref ref-type="bibr" rid="ref204">Simmons 1969</xref>),
<italic>Alternaria</italic> and <italic>Dendryphion</italic>(<xref ref-type="bibr" rid="ref14">Von Arx 1981</xref>). The pluriform
nature of the <italic>Pleospora</italic> anamorphs strongly contrasts with the
relatively uniform morphology of the teleomorphic structures
(<xref ref-type="bibr" rid="ref111">Holm 1962</xref>,
<xref ref-type="bibr" rid="ref126">Kodsueb <italic>et al</italic>. 2006</xref>,
<xref ref-type="bibr" rid="ref116">Inderbitzin <italic>et al</italic>.
2009</xref>). The polyphyletic nature of <italic>Pleospora</italic> has been
hypothesised by Holm (<xref ref-type="bibr" rid="ref111">1962</xref>)
and Berbee (<xref ref-type="bibr" rid="ref27">1996</xref>), but only
recently have molecular studies confirmed its taxonomic complexity
(<xref ref-type="bibr" rid="ref82">Dong <italic>et al</italic>. 1998</xref>,
<xref ref-type="bibr" rid="ref126">Kodsueb <italic>et al</italic>. 2006</xref>,
<xref ref-type="bibr" rid="ref116">Inderbitzin <italic>et al</italic>.
2009</xref>).</p></sec><sec><title>Clade 8, Didymellaceae:</title><p>The major cluster observed in the generic phylogeny is the top clade in
<xref ref-type="fig" rid="fig1">Fig. 1</xref>, which represents the
<italic>Didymellaceae</italic> clade. This clade is well supported (BPP = 0.99, RBS =
94 %), but with the loci used, a high level of basal polytomy is recorded
within the clade. The ancestral species in this clade are the
<italic>Graminae</italic>-pathogens, <italic>Ascochyta hordei</italic> and <italic>Ph.
paspali</italic>. The latter species has been considered to be an indigenous
pathogen of grasses in Australia and New Zealand
(<xref ref-type="bibr" rid="ref119">Johnston 1981</xref>,
<xref ref-type="bibr" rid="ref55">Boerema <italic>et al</italic>. 2004</xref>),
but based on sequence comparisons this species is probably also present in
Europe (<xref ref-type="bibr" rid="ref237">Wirsel <italic>et al</italic>.
2001</xref>, C. Gueidan pers. comm.).</p><p>Clade 8 comprises most <italic>Phoma</italic> species, including
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=615.75&link_type=cbs">CBS 615.75</ext-link>, the
representative strain of <italic>Ph. herbarum</italic>(<xref ref-type="bibr" rid="ref55">Boerema <italic>et al</italic>.
2004</xref>), which is type species of the genus
(<xref ref-type="bibr" rid="ref29">Boerema 1964</xref>). This clade
also includes the type species of the <italic>Phoma</italic> sections <italic>Phoma,
Peyronellaea, Phyllostictoides, Sclerophomella</italic> and <italic>Macrospora</italic>.
Some phytopathologically and medically relevant species of the section
<italic>Heterospora</italic> are also associated with this clade, although some
species of this section are found in other clades, such as <italic>Ph.
samarorum</italic> (clade 6) and <italic>Ph. dimorphospora,</italic> and the sectional
type <italic>Ph. heteromorphospora</italic> (clade 7). Finally, a single species of
the setose section <italic>Paraphoma, Ph. gardeniae</italic>, is found in the
<italic>Didymellaceae</italic>. Based on the sequence data obtained in this study, it
is estimated that approximately 70 % of the species recognised by Boerema
<italic>et al</italic>. (<xref ref-type="bibr" rid="ref55">2004</xref>) can be
associated with the <italic>Didymellaceae</italic>.</p><p>Besides the many <italic>Phoma</italic> species, several other anamorph fungi are
found within this clade, including <italic>Ampelomyces quercinus, Ascochyta
fabae</italic> (teleom<italic>. Didymella fabae</italic>), <italic>Asc. hordei</italic> var.<italic> hordei, Asc. pinodes</italic> (teleom<italic>. Didymella pinodes</italic>),
<italic>Chaetasbolisia erysiphoides, Didymella exigua, Epicoccum nigrum</italic>(synanamorph <italic>Ph. epicoccina</italic>) and <italic>Microsphaeropsis olivacea</italic>.
Of these species, <italic>Asc. pisi, C. erysiphoides</italic> and <italic>M. olivacea</italic>are recognised as type species for their respective genera. De Gruyter <italic>et
al</italic>. (<xref ref-type="bibr" rid="ref100">2009</xref>) already
discussed the probability of finding most of these non-<italic>Phoma</italic> taxa in
the <italic>Didymellaceae</italic> clade.</p><p>It should be noted that not all <italic>Ascochyta</italic> species are found within
this clade, indicating that this genus is also polyphyletic. Whereas <italic>A.
hordei</italic> var. <italic>hordei</italic> is found to be one of the basal taxa of clade
8, the legume associated pathogens <italic>A. fabae, A. pinodes</italic> and the type
species <italic>A. pisi</italic> are found in close association with several species
of <italic>Phoma</italic>. This result is in congruence with the observations in the
study of Peever <italic>et al.</italic>(<xref ref-type="bibr" rid="ref163">2007</xref>). Also the recently
described <italic>Didymella clematidis</italic> has an anamorph state in
<italic>Ascochyta</italic> and is closely related to <italic>Phoma</italic> taxa in this major
clade (<xref ref-type="bibr" rid="ref239">Woudenberg <italic>et al</italic>.
2009</xref>). A representative strain <italic>Asc. caulina</italic> and a new
<italic>Ascochyta</italic> species that is still due to be published (G.J.M. Verkley,
pers. comm.), however, have been found to be only distantly related and are
found in clades 7 and 6, respectively.</p><p>Where a sexual state is known for the <italic>Phoma</italic> and <italic>Ascochyta</italic>species in clade 8, it is <italic>Didymella</italic>. The type species of this
teleomorph genus, <italic>D. exigua</italic>, is also found within this clade,
although it is not associated with a <italic>Phoma</italic> anamorph state. The family
<italic>Didymellaceae</italic> was introduced for this group by De Gruyter <italic>et
al</italic>. (<xref ref-type="bibr" rid="ref100">2009</xref>). However,
type species of two other teleomorph genera have also been found within this
clade. DNA sequences of <italic>Leptosphaerulina australensis</italic> resemble a high
level of similarity with those of the various <italic>Phoma</italic> and
<italic>Didymella</italic> strains, although none are identical. Also sequences of LSU
and ITS sequence data obtained from GenBank of <italic>L. americana, L.
argentinensis, L. chartarum, L. crassiasca</italic> and <italic>L. trifolii</italic>(GenBank accession no. AY278318, AY849949, EU272493, U79485, AY8315585
respectively) were highly similar or even identical to the
<italic>Didymellaceae</italic> sequences obtained in the present study (data not
shown). These observations are in congruence with the results obtained by
Silva-Hanlin & Hanlin
(<xref ref-type="bibr" rid="ref203">1999</xref>), who found that <italic>D.
bryoniae</italic> (anam. <italic>Ph. cucurbitacearum</italic>) was closely related with
<italic>L. chartarum</italic> and <italic>L. crassiasca</italic>. Also <italic>Macroventuria
anomochaeta</italic>, which represents the genus <italic>Macroventuria</italic>(<xref ref-type="bibr" rid="ref1">Van der Aa 1971</xref>) groups in
<italic>Didymellaceae</italic>. The close genetical resemblance of
<italic>Macroventuria</italic> and <italic>Leptosphaerulina</italic> found in the present
study is in congruence with the results of Kodsueb <italic>et al.</italic>(<xref ref-type="bibr" rid="ref126">2006</xref>).</p><p>The loci employed here for phylogenetic analysis are sufficient to identify
clades at the family level, but for proper resolution at generic level or
lower, additional gene regions need to be sequenced. As the majority of
<italic>Phoma</italic> species is embedded in the <italic>Didymellaceae</italic> clade, we
will define further generic and species boundaries within this recently
established family in the subsequent part of this paper.</p></sec></sec></sec><sec><title>Systematics of the <italic>Didymellaceae</italic></title><sec><title>DNA phylogenetic analysis</title><p>The alignment that was used to delineate the <italic>Didymellaceae</italic>consisted of 274 sequences belonging to 196 species. A list of the species
names and numbers, original substrates, geographical origins and GenBank
Accession numbers of the strains used in this study is provided in
<xref ref-type="table" rid="tbl3">Table 3</xref>. The sequence matrix
had a total length of 2 188 characters including the alignment gaps (LSU: 1
327; ITS: 508 and TUB: 353). Of those characters, 1 788 (LSU: 1 233; ITS: 374
and TUB: 181) were constant, whereas 400 characters (LSU: 94; ITS: 144 and
TUB: 192) were variable.</p><p>The analysis run of the LSU-ITS-TUB sequence matrix in MrBayes was aborted
after obtaining 20 000 trees, which was well after stationarity in the
probability of the trees was reached, whereas the standard deviation of split
frequencies was below 0.02. From the obtained tree population, the 25 %
burn-in was discarded and the consensus tree and posterior probabilities were
calculated. The topology and support values of the BI tree were in congruence
with the optimal tree obtained in the ML analysis.</p></sec><sec><title>Systematics: treatment of clades</title><p>As most other anamorph genera, <italic>Phoma</italic> has largely been used as a
convenient form genus, rather than a phylogenetic entity. With the number of
<italic>Phoma</italic> species that are being analysed on DNA sequence level rapidly
increasing, the question is raised whether form genera should be maintained or
that more natural groupings, merging both phylogeny and morphological data,
should be erected. Of course, as greater numbers of taxa are collected and
analysed, the taxonomic boundaries of more clades will be resolved. However,
for the present, only those genera that could be resolved based on available
cultures are treated. The groups mentioned below refer to those indicated
A–R in <xref ref-type="fig" rid="fig2">Fig. 2</xref>. The
unresolved clades are left untreated, and are thus not discussed.</p><p>The taxa in this part of the study were selected based on genetic and/or
morphological similarities with the species that were associated with the
<italic>Didymellaceae</italic> in <xref ref-type="fig" rid="fig1">Fig.
1</xref>. Although numerous taxa from various genera have been associated
with “<italic>Phoma</italic>”, the number of genera that could be included
in the selection for the <italic>Didymellaceae</italic> was limited. Next to
<italic>Phoma</italic>, the only species found were those accommodated on basis of
previous morphological studies in either <italic>Ampelomyces, Ascochyta,
Chaetasbolisia, Coniothyrium, Didymella, Diplodina, Dothiorella, Epicoccum,
Leptosphaerulina, Macroventuria</italic>, or <italic>Microsphaeropsis</italic>. Of three
of these generic representatives, viz. <italic>Chaetasbolisia, Diplodina</italic> and
<italic>Dothiorella</italic>, we suspect that some cultures have been preserved under
an incorrect name. The species representing <italic>Ampelomyces, A.
quercinus,</italic> was correctly identified, but as suggested earlier, the
taxonomic placement in this genus appears to be incorrect
(<xref ref-type="bibr" rid="ref221">Szentiványi <italic>et al.</italic>2005</xref>).</p><p>Strains belonging to a single species proved to be genetically identical or
at least highly similar, indicating that the initial identification of these
strains had been carried out correctly.</p><p>Several well-supported clusters are recognised within this family that are
treated here as novel groups of the <italic>Didymellaceae</italic>. In this section
these separate groups are treated. However, although multiple genes were
employed in this study to generate a phylogenetic reconstruction of the
family, high levels of basal polytomy were observed as well
(<xref ref-type="fig" rid="fig2">Fig. 2</xref>). Application of
general nrDNA loci alone did not reduce this high level of polytomy, whilst
interspecies variation in several well-supported clades was reduced
drastically.</p><sec><title>Group A – outgroup and basal lineages:</title><p>The tree presented in <xref ref-type="fig" rid="fig2">Fig. 2</xref>is rooted to <italic>Ascochyta hordei</italic> and <italic>Ph. paspali</italic>, which proved
to be ancestral to the <italic>Didymellaceae</italic> in
<xref ref-type="fig" rid="fig1">Fig. 1</xref>. The latter species was
described by Johnston (<xref ref-type="bibr" rid="ref119">1981</xref>)
as a species from grasses in New Zealand and Australia, but in recent years,
isolates with similar genotypes were isolated from iron-rich volcanic soil
from France (C. Gueidan, pers. comm.), and from common reed (<italic>Phragmites
australis</italic>) in Germany (<xref ref-type="bibr" rid="ref237">Wirsel <italic>et
al.</italic> 2001</xref>). These isolates were, however, never studied
morphologically.</p><p>Another species used as outgroup is <italic>Ascochyta hordei</italic> var.
<italic>hordei</italic> (<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=544.74&link_type=cbs">CBS
544.74</ext-link>), which was obtained from a South African <italic>Triticum
aestivum</italic>, indicating that also within the <italic>Didymellaceae</italic>, species
that are ascribed to <italic>Ascochyta</italic> do not form a monophyletic group. Also
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=259.92&link_type=cbs">CBS 259.92</ext-link>, the
isotype of <italic>Ph. matteuciicola</italic>, proved to be basal to most other
<italic>Phoma</italic> species. <italic>Phoma matteuciicola</italic> is commonly known as a
pathogen of many fern species (<xref ref-type="bibr" rid="ref99">De Gruyter
<italic>et al.</italic> 2002</xref>). Within the basal lineages, also a group
comprising <italic>Ph. humicola</italic> and the novel species <italic>Ph. saxea</italic> is
found, although this group is only supported by BI analysis (BPP = 0.92, RBS
< 50 %). Although <italic>Phoma humicola</italic> is known as a saprobic soil
fungus, it is sometimes mistaken for the notorious potato pathogen <italic>Ph.
foveata</italic> (Group N), due to a similar biochemical reaction to NaOH and the
formation of citrine green crystals on MEA
(<xref ref-type="bibr" rid="ref98">De Gruyter <italic>et al.</italic>1998</xref>). However, conidia of <italic>Ph. humicola</italic> are always
eguttulate in contrast to those of <italic>Ph. foveata. Phoma saxea</italic> has been
found twice in Germany on rock material, and will be further described
below.</p><p><italic><bold>Phoma humicola</bold></italic> J.C. Gilman & E.V. Abbott, Iowa St.
Coll. J. Sci. 1(3): 266. 1927.</p><p><italic>Specimen examined</italic>: <bold>U.S.A.</bold>, Nevada, Death Valley, from a
dead leaf of <italic>Franseria</italic> sp., 1971, G.H. Boerema, CBS H-16390, culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=220.85&link_type=cbs">CBS 220.85</ext-link>.</p><p><italic><bold>Phoma matteuciicola</bold></italic> Aderkas, Gruyter, Noordel. &
Strongman, Canad. J. Pl. Pathol. 14(3): 227. 1992.</p><p><italic>Specimen examined</italic>: <bold>Canada</bold>, Nova Scotia, Five Mile River,
from leaf base of <italic>Matteuccia struthiopteris</italic>, May 1981, P. von
Aderkas, <bold>holotype</bold> DAOM 183092, culture ex-holotype
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=259.92&link_type=cbs">CBS 259.92</ext-link> = IMI
286996 = PD 91/272.</p><p><italic>Notes</italic>: Gangrene in ostrich fern was originally attributed to
<italic>Ph. exigua</italic> var. <italic>foveata</italic>(<xref ref-type="bibr" rid="ref9">von Aderkas & Brewer
1983</xref>), which is here recombined as <italic>Boeremia foveata</italic>, but
Von Aderkas <italic>et al.</italic>(<xref ref-type="bibr" rid="ref10">1992</xref>) recognised a new
species as causal agent of this disease. The phylogeny presented here supports
these observations, as <italic>Ph. matteuciicola</italic> is found rather distinct
from <italic>B. foveata</italic>.</p><p><italic><bold>Phoma paspali</bold></italic> P.R. Johnst., New Zealand J. Bot. 19(2): 181.
1981.</p><p><italic>Specimens examined</italic>: <bold>New Zealand</bold>, Auckland, Kaikohe, from a
dead leaf of <italic>Paspalum dilatatum</italic>, Jan. 1979, P.K. Buchanan,
<bold>isotype</bold> CBS H-7623, culture ex-isotype
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=560.81&link_type=cbs">CBS 560.81</ext-link> = PD
92/1569; Waikato District, Ruakura, from <italic>Lolium perenne</italic>, Jan. 1979,
G.H. Boerema, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=561.81&link_type=cbs">CBS
561.81</ext-link> = PDDCC 6615.</p><p><italic><bold>Phoma saxea</bold></italic> Aveskamp, Gruyter & Verkley, <bold>sp.
nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515591&link_type=mb">MB515591</ext-link>.
<xref ref-type="fig" rid="fig3">Fig. 3</xref>.</p><p>Conidia dimorpha, intra idem pycnidia formata. Conidia typus 1
(sub)globosa, glabra, hyalina, continua, (3–)3.5–5.5 μm diam.,
(0–)3–10(–15) guttulis praedita. Conidia typus 2 cylindrica
vel ellipsoidea, glabra, hyalina, continua,
(3.5–)4.5–7(–7.5) × 2.5–3.5(–4) μm,
plerumque eguttulata, vel 1–3 guttulis praedita. Matrix conidiorum
salmonea. Chlamydosporae continuae, globosae, viridulae, in catenas usque 35
positae, (8.5–)10–16.5(–17.5) ×
(6–)8–12.5(–14) μm.</p><p><italic>Etymology</italic>: Refers to the substratum on which both isolates of this
species were found, stone material.</p><p><italic>Pycnidia</italic> solitary, (sub-)globose, glabrous or covered with hyphal
outgrows, (90–)135–280(–310) ×
(90–)105–260(–275) μm. <italic>Ostioles</italic> single,
papillate, with wide openings, <italic>ca</italic>. 40–80 μm diam.
<italic>Pycnidial wall</italic> pseudoparenchymatous, composed of oblong to
isodiametric cells, 2–3 layers, 10–17 μm thick, outer cell
layer brown pigmented. <italic>Conidiogenous cells</italic> phialidic, hyaline,
simple, smooth, variable in appearance, flask-shaped, oblong or isodiametric
<italic>ca</italic>. 5.5–7.5 × 3–4 μm. <italic>Conidia</italic> of two
types, both originating from the same pycnidia. Conidia of type 1:
(sub-)globose, thin-walled, smooth, hyaline, aseptate (3–)3.5–5.5
μm diam, with (0–)3–10(–15) guttules. Conidia of type 2:
cylindrical to ellipsoidal, thin-walled, smooth, hyaline, aseptate,
(3.5–)4.5–7(–7.5) × 2.5–3.5(–4) μm,
mainly egutullate or with up to 3 minute guttules. <italic>Conidial matrix</italic>salmon. <italic>Chlamydospores</italic> ubiquitously present in the agar, unicellular,
globose, in long chains of up to 35 elements, greenish pigmented, measuring
(8.5–)10–16.5(–17.5) ×
(6–)8–12.5(–14) μm.</p><p><italic>Culture characteristics</italic>: Colonies on OA, 45–50 mm diam after
7 d, margin regular. Immersed mycelium flat, olivaceous to greenish
olivaceous, citrine-green or coral near the colony margin. Aerial mycelium
absent, but sometimes some grey erect tufts are encountered near the colony
centre; reverse concolourous. Colonies on MEA 20–25 mm diam after 7 d,
margin regular. Immersed mycelium violet-slate, but saffron near the colony
margin. Abundant pycnidia are present on the agar surface; reverse iron-grey,
saffron near the colony margin. Colonies on CHA similar as on MEA, but
somewhat slower growing, 10–15 mm diam. after 7 d, and some sparse white
aerial mycelia hyphae are present in the colony. Application of NaOH results
in a greenish yellow discolouration of the agar, best to be observed on OA
medium.</p><p><italic>Specimens examined</italic>: <bold>Germany</bold>, Oldenburg, from corroded
Mediterranean marble, June 1992, J. Kuroczkin, <bold>holotype designated here</bold>CBS H-20240, culture ex-holotype
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=419.92&link_type=cbs">CBS 419.92</ext-link>;
Oldenburg, from limestone, 1987, J. Kuroczkin,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=298.89&link_type=cbs">CBS 298.89</ext-link>.</p><p><italic>Notes</italic>: The pycnidial wall of <italic>Phoma saxea</italic> is extremely
thin and almost hyaline when the conidia have exuded. Older pycnidia collapse
and remain as a double-layered, disc-like structure on the agar.</p><p>Both strains of this species have been isolated from stone material, such
as limestone (<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=298.89&link_type=cbs">CBS
298.89</ext-link>) and corroded Mediterranean marble
(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=419.92&link_type=cbs">CBS 419.92</ext-link>).
Although the genus is known from all kinds of substrates, the number of
rock-inhabiting <italic>Phoma</italic> isolates is relatively low. Selbmann <italic>et
al.</italic> (<xref ref-type="bibr" rid="ref199">2002</xref>) report on
<italic>Ph. herbarum</italic> from Antarctic rock, and Boerema <italic>et al.</italic>(<xref ref-type="bibr" rid="ref55">2004</xref>) list several species
from rock-like materials, such as cement (<italic>Ph. herbarum</italic>), wall-plaster
(<italic>Ph. heteroderae</italic> – here recombined into <italic>Ph.
calorpreferens</italic>) and crockery (<italic>Ph. pomorum</italic>). In addition,
multiple species are recorded from rock-inhabiting lichens
(<xref ref-type="bibr" rid="ref148">Nelson <italic>et al</italic>. 2009</xref>,
<xref ref-type="bibr" rid="ref186">Ruibal <italic>et al.</italic> 2009</xref>).
These species, listed by Hawksworth & Cole
(<xref ref-type="bibr" rid="ref104">2004</xref>) are, however,
unculturable and could therefore not be compared with <italic>Ph. saxea in
vitro</italic>. However, the morphological descriptions suggest that the mentioned
species and <italic>Ph. saxea</italic> are different taxonomic entities.</p><p><fig position="float" id="fig3"><label>Fig. 3.</label><caption><p><italic>Phoma saxea</italic> (<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=419.92&link_type=cbs">CBS
419.92</ext-link>). A–C. Fourteen-day-old colonies on OA (A), MEA (B)
and CHA (C). D. Pycnidia. E–F. Conidiogenous cells. G. Conidia. H. Chain
of unicellular chlamydospores. Scale bars: D = 100 μm; E–G = 5 μm;
H = 20 μm.</p></caption><graphic xlink:href="1fig3"></graphic></fig></p></sec><sec><title>Group B:</title><p>Four of the six species clustering in Group B produce a <italic>Didymella</italic>teleomorph. Only <italic>Ph. polemonii</italic> and <italic>Ph. xanthina</italic> presently
have no known sexual state. The species in this clade are collected from a
wide variety of dicots, although all individual taxa appear to be
host-specific (<xref ref-type="bibr" rid="ref55">Boerema <italic>et al.</italic>2004</xref>). Also the micromorphological features of these species are
highly variable.</p><p>A single strain that was kept in the CBS collection as <italic>Diplodina
coloradensis</italic> was found in this clade as well. However, this genus name
has been accommodated in the <italic>Gnomoniaceae</italic> (<italic>Diaporthales</italic>),
indicating that this strain has been preserved under an incorrect name and
should be renamed. However, as this strain proved to be sterile, no proper
redescription of the material could be provided.</p><p><italic><bold>Didymella applanata</bold></italic> (Niessl) Sacc., Syll. Fung. 1: 546.
1882.</p><p><italic>Basionym</italic>: <italic>Didymosphaeria applanata</italic> Niessl, Oesterr. Bot.
Z. 25(4): 129. 1875.</p><p><italic>Anamorph</italic>: <italic><bold>Phoma argillacea</bold></italic> (Bres.) Aa &
Boerema, in De Gruyter, Boerema & Van der Aa, Persoonia 18(1): 17.
2002.</p><p><italic>Basionym</italic>: <italic>Phyllosticta argillacea</italic> (Bres.), Hedwigia 1894:
206. 1894.</p><p><italic>Specimens examined</italic>: <bold>The Netherlands</bold>, Baarn, from <italic>Rubus
idaeus</italic>, Sep. 1963, A. van Dijkman, CBS H-11943, culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=205.63&link_type=cbs">CBS 205.63</ext-link>; from
<italic>Rubus idaeus</italic>, 1975, G.H. Boerema,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=102634&link_type=cbs">CBS 102634</ext-link> = PD
75/248.</p><p><italic><bold>Didymella cannabis</bold></italic> (G. Winter) Arx, in Müller &
Arx, Beitr. Kryptogamenfl. Schweiz 11(2): 365. 1962.</p><p><italic>Basionym</italic>: <italic>Sphaerella cannabis</italic> G. Winter, Hedwigia 11(10):
145. 1872.</p><p><italic>Anamorph. <bold>Phoma cannabis</bold></italic> (L.A. Kirchn.) McPartl., Mycologia
86(6): 871. 1995.</p><p><italic>Basionym</italic>: <italic>Depazea cannabis</italic> L.A. Kirchn., Lotos 6: 183.
1856.</p><p><italic>Specimen examined</italic>: Unknown origin, from <italic>Cannabis sativa</italic>,
Oct. 1937, K. Röder, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=234.37&link_type=cbs">CBS
234.37</ext-link>.</p><p><italic>Notes</italic>: The studied culture
(<xref ref-type="bibr" rid="ref183">Röder 1937</xref>) is now
sterile, and could therefore not be described here morphologically.</p><p><italic><bold>Didymella catariae</bold></italic> (Cooke & Ellis) Sacc., Syll. Fung.
1: 557. 1882.</p><p><italic>Basionym</italic>: <italic>Sphaeria catariae</italic> Cooke & Ellis, Grevillea
5: 96. 1876.</p><p><italic>Anamorph</italic>: <italic><bold>Phoma nepeticola</bold></italic> (Melnik) Dorenb. &
Gruyter, Persoonia 18(1): 18. 2002.</p><p><italic>Basionym</italic>: <italic>Ascochyta nepeticola</italic> Melnik, Novoste Sist.
Nizsh. Rast. 1968: 178. 1968.</p><p><italic>Specimen examined</italic>: <bold>The Netherlands</bold>, from the stem of
<italic>Nepeta catenaria</italic>, 1977, M.M.J Dorenbosch,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=102635&link_type=cbs">CBS 102635</ext-link> = PD
77/1131.</p><p><italic><bold>Didymella urticicola</bold></italic> Aa & Boerema, in Boerema, Trans.
Brit. Mycol. Soc. 67(2): 303. 1976.</p><p><italic>Anamorph</italic>: <italic><bold>Phoma urticicola</bold></italic> Aa & Boerema, in
Boerema, Trans. Brit. Mycol. Soc. 67(2): 303. 1976.</p><p><italic>Specimens examined</italic>: <bold>The Netherlands</bold>, Wageningen, from a
dead stem tip of <italic>Urtica dioica</italic>, Mar. 1973, G.H. Boerema, holotype CBS
H-11971, culture ex-holotype
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=121.75&link_type=cbs">CBS 121.75</ext-link> = ATCC
32164 = IHEM 3403 = IMI 194767 = PD 73/584; from <italic>Urtica dioica</italic>, 1973,
G.H. Boerema, PD 73/570.</p><p><italic><bold>Phoma polemonii</bold></italic> Cooke, Grevillea 13(68): 94. 1885.</p><p><italic>Specimen examined</italic>: <bold>The Netherlands</bold>, from <italic>Polemonium
caeruleum</italic>, 1983, J. de Gruyter,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=109181&link_type=cbs">CBS 109181</ext-link> = PD
83/757.</p><p><italic><bold>Phoma xanthina</bold></italic> Sacc., Michelia 1(4): 359. 1884.</p><p><italic>Specimens examined</italic>: <bold>The Netherlands</bold>, Baarn, from leaves of
<italic>Delphinium</italic> sp., May 1968, H.A. van der Aa, CBS H-8938, culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=383.68&link_type=cbs">CBS 383.68</ext-link>; from
<italic>Delphinium</italic> sp., 1984, G.H. Boerema, PD 84/407.</p></sec><sec><title>Group C:</title><p>The species in Group C cluster in two subgroups: One comprising the
<italic>Clematis</italic> pathogens <italic>Ph. clematidina</italic> and <italic>Coniothyrium
clematidis-rectae</italic>, the other subgroup comprising <italic>Ph.
aquilegiicola</italic> and <italic>Ph. glaucii</italic>, two pathogens of
<italic>Ranunculaceae</italic> and <italic>Papaveraceae,</italic> respectively. All three
<italic>Phoma</italic> species in this group were morphologically linked to the
section <italic>Heterospora</italic> (<xref ref-type="bibr" rid="ref54">Boerema
<italic>et al.</italic> 1997</xref>), but are distinct from the species in clade
S by the absence of conidia that represent the <italic>Stagonosporopsis</italic>synanamorph in culture, although smaller septate conidia do occur. In these
species the <italic>Stagonosporopsis</italic> synanamorph is only known from <italic>in
vivo</italic> material (<xref ref-type="bibr" rid="ref33">Boerema
1993</xref>, <xref ref-type="bibr" rid="ref54">Boerema <italic>et al</italic>.
1997</xref>).</p><p>The several species that were associated with the <italic>Ph. clematidina</italic>morphotype have recently been distinguished in a study of Woudenberg <italic>et
al.</italic> (<xref ref-type="bibr" rid="ref239">2009</xref>). In the same
study, the authors showed that <italic>C. clematidis-rectae</italic> is closely
related and, based on sequence analysis, a member of the family
<italic>Didymellaceae</italic>. The major character on which this species is regarded
as distinct from <italic>Ph. clematidina</italic> is by the production of pale brown
pigmented conidia. In addition, the conidiogenesis of <italic>Coniothyrium</italic> is
annellidic with percurrent proliferation, in contrast to the conidiogenesis in
<italic>Phoma</italic>, which is considered to be solely phialidic with periclinal
thickening (<xref ref-type="bibr" rid="ref36">Boerema & Bollen
1975</xref>, <xref ref-type="bibr" rid="ref219">Sutton
1980</xref>). Evidence for the presence of annellides has, however, not
been observed in <italic>C. clematidis-rectae</italic>, while conidial pigmentation is
relatively pale in comparison to other <italic>Coniothyrium</italic> species.
Pigmented conidia have also been observed in various <italic>Phoma</italic> species
before (<xref ref-type="bibr" rid="ref83">Dorenbosch 1970</xref>,
<xref ref-type="bibr" rid="ref55">Boerema <italic>et al.</italic> 2004</xref>,
<xref ref-type="bibr" rid="ref19">Aveskamp <italic>et al.</italic>2009a</xref>). These features may indicate that this species is actually
a <italic>Phoma</italic> with early conidial pigmentation. Therefore <italic>C.
clematidis-rectae</italic> is recombined into <italic>Phoma</italic> below.</p><p><italic><bold>Phoma aquilegiicola</bold></italic> M. Petrov, Acta Inst. Bot. Acad. Sci.
USSR Pl. Crypt. [Trudy Bot. Inst. Akad. Nauk SSSR] Fasc. 1: 281. 1933.</p><p><italic>Specimens examined</italic>: <bold>The Netherlands</bold>, from a stem of
<italic>Aconitum pyramidale</italic>, 1973, G.H. Boerema,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=107.96&link_type=cbs">CBS 107.96</ext-link> = PD
73/598; from a stem of <italic>Aquilegia</italic> sp., 1979, G.H. Boerema,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=108.96&link_type=cbs">CBS 108.96</ext-link> = PD
79/611.</p><p><italic><bold>Phoma clematidina</bold></italic> (Thüm.) Boerema, Verslagen Meded.
Plziektenk. Dienst Wageningen (Jaarboek 1978) 153: 17. 1979. emend. Woudenberg
<italic>et al</italic>., Persoonia 22: 59. 2009.</p><p><italic>Basionym</italic>: <italic>Ascochyta clematidina</italic> Thüm., Bull. Soc.
Imp. Naturalistes Moscou 55: 98. 1880.</p><p><italic>Specimens examined</italic>: <bold>Russia</bold>, Minussinsk, from leaves of
<italic>Clematis glaucae</italic>, N. Martianoff, <bold>isotype</bold> LE 40082; <bold>The
Netherlands</bold>, Spaubeek, from the stem of <italic>Clematis</italic> sp., July 1978,
<italic>G.H. Boerema</italic>, <bold>epitype</bold> CBS H-16193, culture exepitype
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=108.79&link_type=cbs">CBS 108.79</ext-link> = PD
78/522; from <italic>Clematis</italic> sp., I. de Boer, Nov. 1949,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=201.49&link_type=cbs">CBS 201.49</ext-link>;
Boskoop, from <italic>Clematis jackmanii</italic>, C. Dorsman, Oct. 1962,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=195.64&link_type=cbs">CBS 195.64</ext-link>;
Wageningen, from <italic>Selaginella</italic> sp. M.M.J. Dorenbosch, 1966,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=520.66&link_type=cbs">CBS 520.66</ext-link>;
<bold>U.K.</bold>, England, from <italic>Clematis</italic> sp., Jan. 1966, F.T. Last,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=102.66&link_type=cbs">CBS 102.66</ext-link>.</p><p><italic><bold>Phoma clematidis-rectae</bold></italic> (Petr.) Aveskamp, Woudenberg &
Gruyter, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515592&link_type=mb">MB515592</ext-link>.</p><p><italic>Basionym</italic>: <italic>Coniothyrium clematidis-rectae</italic> Petr., Fungi
Polon. 576. 1921.</p><p><italic>Pycnidia</italic> solitary or confluent, immersed or produced on the agar
surface, globose, glabrous, (80–)85–130(–155) μm diam, in
older cultures pycnidia may become larger and grow after maturation to
220–250 μm diam. <italic>Ostioles</italic> 1(–4), wide, non-papillate
to papillate or, in older cultures, on a elongated neck. <italic>Pycnidial
wall</italic> pseudoparenchymatous, composed of oblong to isodiametric cells,
4–5 layers, (10–)11–19(–19.5) μm thick, outer
1–2 layers pigmented. <italic>Conidiogenous cells</italic> phialidic, hyaline,
simple, smooth, ampulliform to doliiform, measuring 3–4.5(–5)
× 2.5–4.5 μm. <italic>Conidia</italic> ellipsoidal to cylindrical,
thin-walled, smooth, aseptate, (3–) 4–7(–8) ×
2–3(–3.5) μm, with (2–)5–12 guttules, initially
hyaline, but mature conidia become slightly brownish pigmented. <italic>Conidial
matrix</italic> sepia.</p><p><italic>Culture characteristics</italic>: Colonies on OA 42–52 mm diam after
7 d, margin regular. Immersed mycelium dark brick to sepia or iron-grey, but
hyaline near the colony margin. Pycnidia in concentric rings give the colony
an olivaceous tinge. Aerial mycelium absent; reverse concolourous. Colonies on
MEA 27–52 mm diam after 7 d, margin regular. Aerial mycelium
incidentally occurs in sectors in some strains, grey to olivaceous. Immersed
mycelium rosy-buff to rosy-vinaceous with olivaceous and grey tinges; reverse
olivaceous iron-grey to saffron. Application of NaOH did not have any
effect.</p><p><italic>Specimens examined</italic>: <bold>The Netherlands</bold>, Boskoop, from
<italic>Clematis</italic> sp., 1963, G.H. Boerema, CBS H-20275, culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=507.63&link_type=cbs">CBS 507.63</ext-link> = PD
07/03486747 = MUCL 9574; from <italic>Clematis</italic> sp., 1995, J. de Gruyter, PD
95/1958.</p><p><italic>Notes</italic>: In congruence with the studies of Woudenberg <italic>et
al.</italic> (<xref ref-type="bibr" rid="ref239">2009</xref>), this species
was found to be closely related to <italic>Ph. clematidina</italic> and other
<italic>Didymellaceae</italic> species. In contrast, it is only distantly related to
the type species of <italic>Coniothyrium, C. palmarum</italic>. Therefore, a
recombination into <italic>Phoma</italic> is proposed here. The present species is
clearly distinct from <italic>Ph. clematidina</italic> by the production of pigmented
conidia, although the level of pigmentation is low, which distinguishes
<italic>Ph. clematidis-rectae</italic> from the species remaining in
<italic>Coniothyrium</italic> that produce darker, olivaceous conidia.</p><p><italic><bold>Phoma glaucii</bold></italic> Brunaud, “<italic>Ph. glauci</italic>”,
Ann. Soc. Sci. Nat. La Rochelle 1892: 97. 1892.</p><p><italic>Specimens examined</italic>: <bold>The Netherlands</bold>, near Lisse, from
<italic>Dicentra</italic> sp., 1979, G.H. Boerema,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=112.96&link_type=cbs">CBS 112.96</ext-link>;
Wageningen, from a leaf of <italic>Chelidonium majus</italic>, 1994, G.H. Boerema,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=114.96&link_type=cbs">CBS 114.96</ext-link> = PD
94/888.</p></sec><sec><title>Groups D & E – Leptosphaerulina and Macroventuria:</title><p>The most remarkable findings in the <italic>Didymellaceae</italic> are the
<italic>Leptosphaerulina</italic> and <italic>Macroventuria</italic> (clade E) teleomorph
genera. The species belonging to these teleomorphs are found amidst the
<italic>Didymellaceae</italic>, causing the genus <italic>Didymella</italic> to be
paraphyletic. The species in both genera are closely related to each other, as
was already pointed out by Kodsueb <italic>et al.</italic>(<xref ref-type="bibr" rid="ref126">2006</xref>), who, however, missed
the link with <italic>Didymella</italic>. A phomoid anamorph state has, thus far, not
been recorded for any of the species in these teleomorph genera.</p><p><italic>Leptosphaerulina</italic> is morphologically distinct from
<italic>Macroventuria</italic> and <italic>Didymella</italic>, although all three genera are
known for their hyaline ascospores (<xref ref-type="bibr" rid="ref1">Van der
Aa 1971</xref>, <xref ref-type="bibr" rid="ref14">Von Arx
1981</xref>). <italic>Leptosphaerulina</italic> produces large, longitudinally
and transversally septated ascospores, resembling those of <italic>Pleospora</italic>and <italic>Cucurbitaria</italic>, although the ascospores of these genera are
pigmented. The major difference between <italic>Didymella</italic> and
<italic>Macroventuria</italic> is the presence of setae on the pseudothecia of the
latter genus, whereas <italic>Didymella</italic> ascomata are commonly glabrous.
According to the original description (<xref ref-type="bibr" rid="ref1">Van
der Aa 1971</xref>), <italic>Macroventuria</italic> strains resemble
<italic>Venturia</italic> by their setose pycnidia, but differ in their restricted
number of the asci.</p><p><italic><bold>Leptosphaerulina americana</bold></italic> (Ellis & Everh.) J.H. Graham
& Luttr., Phytopathology 51: 686. 1961.</p><p><italic>Basionym</italic>: <italic>Pleospora americana</italic> Ellis & Everh., in
North American Pyrenomycetes: 336. 1892, nom. nov. pro <italic>Pleospora
hyalospora</italic> Ellis & Everh., Proc. Acad. Nat. Sci. Philadelphia: 238.
1890, non <italic>Pleospora hyalospora</italic> Speg.</p><p><italic>Specimen examined</italic>: <bold>U.S.A.</bold>, Georgia, from <italic>Trifolium
pratense</italic>, Apr. 1954, E.S. Luttrell,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=213.55&link_type=cbs">CBS 213.55</ext-link>.</p><p><italic><bold>Leptosphaerulina arachidicola</bold></italic> W.Y. Yen, M.J. Chen &
K.T. Huang, J. Agric. Forest. 10: 167. 1956.</p><p><italic>Specimen examined</italic>: <bold>Taiwan</bold>, from a leaf of <italic>Arachis
hypogaea</italic>, 1956, K.T. Huang,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=275.59&link_type=cbs">CBS 275.59</ext-link> = ATCC
13446.</p><p><italic>Note</italic>: <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=275.59&link_type=cbs">CBS
275.59</ext-link> is degenerated and forms only very tiny sclerotia <italic>in
vitro.</italic></p><p><italic><bold>Leptosphaerulina australis</bold></italic> McAlpine, Fungus Diseases of
stone-fruit trees in Australia: 103. 1902.</p><p><italic>Specimens examined</italic>: <bold>Indonesia</bold>, Lampung, from <italic>Eugenia
aromatica</italic>, Dec. 1982, H. Vermeulen,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=317.83&link_type=cbs">CBS 317.83</ext-link>. <bold>The
Netherlands</bold>, Baarn, from soil, Sep. 1969, J.A. Stalpers,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=939.69&link_type=cbs">CBS 939.69</ext-link>.</p><p><italic><bold>Leptosphaerulina trifolii</bold></italic> (Rostr.) Petr., Sydowia 13: 76.
1959.</p><p><italic>Basionym</italic>: <italic>Sphaerulina trifolii</italic> Rostr., Bot. Tidsskr. 22:
265. 1899.</p><p><italic>Specimen examined</italic>: <bold>The Netherlands</bold>, from <italic>Trifolium</italic>sp., 1958, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=235.58&link_type=cbs">CBS
235.58</ext-link>.</p><p><italic><bold>Macroventuria anamochaeta</bold></italic> Aa, Persoonia 6(3): 362.
1971.</p><p><italic>Specimens examined</italic>: <bold>South Africa</bold>, Karroo Desert, from
decayed canvas, Aug. 1971, M.C. Papendorf, <bold>holotype</bold> CBS H-14192,
ex-holotype culture <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=525.71&link_type=cbs">CBS
525.71</ext-link>; Cape Province, from a trunk of <italic>Medicago sativa</italic>,
June 1972, W.F.O. Marasas, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=502.72&link_type=cbs">CBS
502.72</ext-link>.</p><p><italic><bold>Macroventuria wentii</bold></italic> Aa, Persoonia 6(3): 361. 1971.</p><p><italic>Specimen examined</italic>: <bold>U.S.A.</bold>, Nevada, Death Valley, from plant
litter, Aug. 1971, F.W. Went, <bold>holotype</bold> CBS H-14195, ex-holotype culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=526.71&link_type=cbs">CBS 526.71</ext-link>.</p></sec><sec><title>Group F:</title><p>As a sister group to <italic>Leptosphaerulina</italic>, several host-specific
<italic>Phoma</italic>species are found that induce leaf spots on a variety of plant
species, including <italic>Ph. infossa, Ph. anigozanthi, Ph.
arachidis-hypogaeae</italic> and <italic>Ph. gossypiicola</italic>. The latter species
causes leaf spots and stem canker on cotton plants (<italic>Gossypium</italic> spp.).
However, other plant species may also become symptomatic when deliberately
infected (<xref ref-type="bibr" rid="ref110">Holliday & Punithalingam
1970</xref>). <italic>Phoma infossa</italic> has originally been reported from
stems of ash trees (<italic>Fraxinus</italic> sp.) in New York State
(<xref ref-type="bibr" rid="ref85">Ellis & Everhart 1888</xref>),
but has recently been associated with a severe foliar disease of green ash
(<italic>F. pennsylvanica</italic>) in Argentina
(<xref ref-type="bibr" rid="ref19">Aveskamp <italic>et al.</italic>2009a</xref>). All species produce aseptate conidia in culture, although
<italic>Ph. gossypiicola</italic> is known to also produce 2- to multi-celled conidia
<italic>in vivo</italic>, hence the <italic>Ascochyta gossypii</italic> synonym
(<xref ref-type="bibr" rid="ref94">De Gruyter 2002</xref>).</p><p>In contrast to these plant pathogens, a fungicolous species also occurs in
the present clade. Species from the genera <italic>Phoma</italic> and
<italic>Ampelomyces</italic> have been “frequently confused with each
other” (<xref ref-type="bibr" rid="ref214">Sullivan & White
2000</xref>), which explains why <italic>Ph. fungicola</italic> is found here.
This species was previously known as <italic>Amp. quercinus</italic> and is recombined
in the subsequent taxonomical section. The finding of this species in the
<italic>Didymellaceae</italic> is in congruence with sequence results obtained by
Sullivan & White (<xref ref-type="bibr" rid="ref214">2000</xref>)
and Szentiványi <italic>et al.</italic>(<xref ref-type="bibr" rid="ref221">2005</xref>). Also <italic>Amp.
humuli</italic>, another fast-growing species, proved to be phylogenetically
similar to species that currently represent the <italic>Didymellaceae</italic>(<xref ref-type="bibr" rid="ref124">Kiss & Nakasone 1998</xref>).
Additionally, it has been suggested that the fast growing species <italic>Amp.
artemisiae</italic> and <italic>Amp. uncinulae</italic>(<xref ref-type="bibr" rid="ref185">Rudakov 1979</xref>,
<xref ref-type="bibr" rid="ref123">Kiss 1997</xref>) actually do, in
fact, not represent <italic>Ampelomyces</italic>, but belong to the genus
<italic>Phoma</italic>; these species were incorrectly identified based on their
host-association (<xref ref-type="bibr" rid="ref125">Kiss <italic>et al.</italic>2004</xref>). The species in <italic>Ampelomyces</italic> are all recognised as
parasites of fungi that cause powdery mildew
(<xref ref-type="bibr" rid="ref123">Kiss 1997</xref>). However, it is
suggested that also the ubiquitous species <italic>Ph. glomerata</italic> has
fungicolous capacities, and may be suited as mycoparasitic control agent of
powdery mildew (<xref ref-type="bibr" rid="ref214">Sullivan & White
2000</xref>).</p><p>Only one of the <italic>Phoma</italic> species embedded in this clade has been
associated with a teleomorph. In the description of <italic>Ph. anigozanthi</italic>,
the sexual state is recorded as <italic>Sphaerella millepunctata</italic> (apud
<xref ref-type="bibr" rid="ref96">Gruyter & Noordeloos
1992</xref>). <italic>Sphaerella</italic> is practically synonymised with
<italic>Mycosphaerella</italic> (<italic>e.g.</italic><xref ref-type="bibr" rid="ref11">Aptroot 2006</xref>), but as
described above, several of the <italic>Mycosphaeralla</italic> species have
subsequently been recombined into <italic>Didymella</italic>. In the present study no
evidence of teleomorph formation <italic>in vitro</italic> has been observed, which is
in congruence with the results of Gruyter & Noordeloos
(<xref ref-type="bibr" rid="ref96">1992</xref>). As also type material
of <italic>Ph. anigozanthi</italic> and <italic>S. multipunctata</italic> could not be
obtained, this taxonomic link is still to be confirmed.</p><p><italic><bold>Phoma anigozanthi</bold></italic> Tassi, Boll. Reale Orto Bot. Siena 3 (2
– 1899): 148. 1900.</p><p><italic>Specimen examined</italic>: <bold>The Netherlands</bold>, from a leaf of
<italic>Anigozanthus maugleisii</italic>, 1979, H. Cevat CBS H-5199, culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=381.91&link_type=cbs">CBS 381.91</ext-link> = PD
79/1110.</p><p><italic><bold>Phoma arachidis-hypogaeae</bold></italic> (V.G. Rao) Aa & Boerema,
Persoonia 15(3): 388. 1993.</p><p><italic>Basionym</italic>: <italic>Phyllosticta arachidis-hypogaeae</italic> V.G. Rao,
Sydowia 16 (1962): 275. 1963.</p><p><italic>Specimen examined</italic>: <bold>India</bold>, Madras, from a leaf of
<italic>Arachis hypogaea</italic>, 1977,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=125.93&link_type=cbs">CBS 125.93</ext-link> = PD
77/1029.</p><p><italic><bold>Phoma fungicola</bold></italic> Aveskamp, Gruyter & Verkley, <bold>nom.
nov.</bold> pro <italic>Cicinobolus quercinus</italic> Syd. Ann. Mycol. 13: 42. 1915.
MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515593&link_type=mb">MB515593</ext-link>.</p><p><italic>Basionym</italic>: <italic>Cicinobolus quercinus</italic> Syd., Ann. Mycol. 13: 42.
1915.</p><p><list list-type="simple"><list-item><p>≡ <italic>Ampelomyces quercinus</italic> (Syd.) Rudakov, Mikol. Fitopatol.
13(2): 109. 1979. not <italic>Phoma quercina</italic> Sacc & Roum. Syll. fung. 3:
96. 1881, = <italic>Phomopsis quercina</italic> Sacc.) Höhn., not <italic>Phoma
quercina (</italic>Peck) Sacc. Syll. fung. 3: 96. 1884.</p></list-item></list></p><p><italic>Etymology</italic>: Epithet refers to the fungicolous lifestyle of this
species.</p><p><italic>Pycnidia</italic> always solitary, produced on the agar surface, globose,
peroblate to suboblate, glabrous, measuring
(50–)65–130(–150) ×
(65–)95–200(–220) μm with a single, conspicuous,
non-papillate ostiole. <italic>Pycnidial wall</italic> pale brown,
pseudoparenchymatous, composed of isodiametric cells, 3–5 layers,
(6–)8.5–14.5(–16) μm thick, outer 1–2 layers
slightly pigmented. <italic>Conidiogenous cells</italic> phialidic, hyaline, simple,
smooth, doliiform to ampulliform, variable in size, <italic>ca</italic>.
(3–)3.5–5 × 3–4(–5) μm. <italic>Conidia</italic>variable in shape and size, subglobose to oval or obtuse, thin-walled, smooth,
aseptate, measuring (5–)5.5–7.5(–8.5) ×
3–4.5(–5) μm, with 0–2(–3) minute guttules,
initially hyaline, but brown at maturity. Conidial exudates not recorded.</p><p><italic>Culture characteristics</italic>: Colonies on OA 55–68 mm diam after
7 d, margin regular. Aerial mycelium white, floccose to woolly. Immersed
mycelium greenish olivaceous to olivaceous near the colony centre. Abundant
black pycnidia are scattered over the medium; reverse concolourous. Colonies
on MEA 65–75 mm diam after 7 d, margin regular. Aerial mycelium covering
the whole colony, compact, white to pale grey, with olivaceous tinges near the
colony centre; reverse olivaceous-black.</p><p><italic>Specimen examined</italic>: <bold>Ukraine</bold>, Crimea, in the vicinity of
Feodosiya, on <italic>Microsphaera alphitoides</italic> from <italic>Quercus</italic> sp.,
1979, O.L. Rudakov, CBS H-20276, culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=633.92&link_type=cbs">CBS 633.92</ext-link> = ATCC
36786, VKM MF-325.</p><p><italic>Notes</italic>: The epithet used for the description of this species in the
genera <italic>Cicinobolus</italic> and <italic>Ampelomyces</italic> could not be transferred
to the genus <italic>Phoma</italic> as <italic>Ph. quercina</italic> is already occupied. This
name, however, refers to a <italic>Phyllosticta</italic> species
(<xref ref-type="bibr" rid="ref5">Van der Aa & Vanev 2002</xref>).
Therefore, a new name is proposed here for the present species.</p><p>Kiss & Nakasone (<xref ref-type="bibr" rid="ref124">1998</xref>)
already found that several fast-growing <italic>Ampelomyces</italic> species were
phylogenetically distinct from the type species, which is characterised by a
rather slow growth rate, and suggested that <italic>A. quercinus</italic> belonged to
<italic>Phoma</italic>. This finding was supported by results obtained in later
studies (<xref ref-type="bibr" rid="ref214">Sullivan & White
2000</xref>, <xref ref-type="bibr" rid="ref221">Szentiványi
<italic>et al.</italic> 2005</xref>).</p><p><italic><bold>Phoma gossypiicola</bold></italic> Gruyter, Persoonia 18(1): 96. 2002.</p><p><italic>Specimen examined</italic>: <bold>U.S.A.</bold>, Texas, from a leaf of
<italic>Gossypium</italic> sp., 1963, L.S. Bird CBS H-9006, culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=377.67&link_type=cbs">CBS 377.67</ext-link>.</p><p><italic><bold>Phoma infossa</bold></italic> Ellis & Everh., J. Mycol. 4(10): 102.
1888, emend. Aveskamp <italic>et al.</italic>, Mycologia 101. 373. 2009.</p><p><italic>Specimens examined</italic>: <bold>Argentina</bold>, Buenos Aires Province, La
Plata, from leafs of <italic>Fraxinus pennsylvanica</italic>, 2008, M. Murace,
<bold>neotype</bold> CBS H-20145, culture ex-neotype
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123395&link_type=cbs">CBS 123395</ext-link>; Buenos
Aires Province, La Plata, from leafs of <italic>Fraxinus pennsylvanica</italic>, 2008,
M. Murace, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123394&link_type=cbs">CBS
123394</ext-link>.</p></sec><sec><title>Group G:</title><p>This group (BPP = 1.00, RBS = 99 %) consists of <italic>Ph. subherbarum</italic>and <italic>Ph. pedeiae</italic> sp. nov. Although the first species name suggests a
close resemblance with the type species <italic>Ph. herbarum</italic>, it is
phylogenetically distinct. Both <italic>Ph. herbarum</italic> and <italic>Ph.
subherbarum</italic> are accommodated in section <italic>Phoma</italic>, but are distinct
in colony characters: in contrast to <italic>Ph. herbarum, Ph. subherbarum</italic>does not react to the application of a droplet of NAOH
(<xref ref-type="bibr" rid="ref97">De Gruyter <italic>et al.</italic>1993</xref>). The growth rate of <italic>Ph. subherbarum</italic> is also
considerably faster, as a colony can cover the plate surface within 1 wk.</p><p>Boerema <italic>et al.</italic>(<xref ref-type="bibr" rid="ref55">2004</xref>) hypothesised that
<italic>Ph. subherbarum</italic> is from American origin. In contrast, both strains of
<italic>Ph. pedeiae</italic> were found in the Netherlands. Both species in this clade
appear to have a plurivorous nature. The novel species <italic>Ph. pedeiae</italic> is
described below.</p><p><italic><bold>Phoma pedeiae</bold></italic> Aveskamp, Gruyter & Verkley, <bold>sp.
nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515594&link_type=mb">MB515594</ext-link>.
<xref ref-type="fig" rid="fig4">Fig. 4</xref>.</p><p>Conidia ellipsoidea vel cylindrica, glabra, hyalina, continua, 3–4.5
× 1.5–2.5 μm, 0–2(–3) guttulis praedita. Matrix
conidiorum cremeo-alba.</p><p><italic>Etymology</italic>: Named after the institute that has facilitated most of
the research on the taxonomy of the genus <italic>Phoma</italic> and affiliated genera
in the past decade, the PD (Plantenziektenkundige Dienst – Dutch Plant
Protection Service). Both isolates of this species were collected and
preserved by employees of this institute.</p><p><italic>Pycnidia</italic> solitary or confluent, produced on the agar surface,
globose to ellipsoidal, glabrous, (90–)100–230(–255) ×
(75–) 90–155(–165) μm with 1–2 conspicuous,
non-papillate ostioles. <italic>Pycnidial wall</italic> pseudoparenchymatous, composed
of oblong to isodiametric cells, 3–5 layers, 11–17 μm thick.
<italic>Conidiogenous cells</italic> phialidic, hyaline, simple, smooth, flask-shaped,
relatively small, <italic>ca.</italic> 3.5–4(–4.5) × 3–4
μm. <italic>Conidia</italic> ellipsoidal to cylindrical, thin-walled, smooth,
hyaline, aseptate 3–4.5 × 1.5–2.5 μm, with
0–2(–3) guttules. <italic>Conidial matrix</italic> creme-white.</p><p><italic>Culture characteristics</italic>: Colonies on OA, 65–75 mm diam after
7 d, margin regular. Immersed mycelium olivaceous. Aerial mycelium floccose,
white or smoke-grey to greenish olivaceous. Abundant black pycnidia are
scattered over the medium; reverse concolourous with some reddish tinges.
Colonies on MEA 55–65 mm diam after 7 d, margin regular. Aerial mycelium
covering the whole colony, floccose, smoke-grey to greenish olivaceous, white
near the centre of the colony; reverse olivaceous-black or bay. Agar colour
changes to bay due to diffusible pigments produced by the fungus. Colonies on
CHA similar as on MEA, but somewhat faster growing, 70–80 mm diam after
7 d. Application of NaOH did not have any effect.</p><p><italic>Specimens examined</italic>: <bold>The Netherlands</bold>, Aalsmeer region, on
<italic>Schefflera elegantissima</italic>, 1992, J. de Gruyter, <bold>holotype designated
here</bold> CBS H-20239, culture ex-holotype
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=124517&link_type=cbs">CBS 124517</ext-link> = PD
92/612A; on <italic>Orchidaceae</italic> sp., 1984, J. de Gruyter,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=124516&link_type=cbs">CBS 124516</ext-link> = PD
84/453.</p><p><italic>Notes</italic>: <italic>Phoma pedeiae</italic> has been found in association with
several tropical ornamental pot plants in Dutch greenhouses. Only mild disease
symptoms were recorded from this species, and therefore the fungus was not
further studied. Phylogenetically, this species is found in close relation to
<italic>Ph. subherbarum</italic> (BPP= 1.00; RBS = 99 %), which is probably a weak
pathogen and saprobe of different plant substrates occurring on the American
continent (<xref ref-type="bibr" rid="ref97">De Gruyter <italic>et al.</italic>1993</xref>).</p><p><italic><bold>Phoma subherbarum</bold></italic> Gruyter, Noordel. & Boerema,
Persoonia 15(3): 387. 1993.</p><p><italic>Specimens examined</italic>: <bold>Canada</bold>, from <italic>Zea mays</italic>,
<bold>holotype</bold> L 992.177.439, culture ex-holotype
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=250.9292&link_type=cbs">CBS 250.9292</ext-link> =
DAOM 171914 = PD 92/371; from <italic>Zea mays</italic>, May 1978, G.A. Neish,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=305.79&link_type=cbs">CBS 305.79</ext-link>A = DAOM
170848; <bold>Peru</bold>, from <italic>Solanum</italic> sp.,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=249.92&link_type=cbs">CBS 249.92</ext-link> = PD
78/1088.</p></sec><sec><title>Group H:</title><p><italic>Phoma bellidis</italic> and <italic>Ph. senecionis</italic> are found in
association with two plant genera from the <italic>Compositae</italic> family:
<italic>Bellis</italic> spp. and <italic>Senecio</italic> spp. respectively
(<xref ref-type="bibr" rid="ref97">De Gruyter <italic>et al.</italic>1993</xref>). The distantly related <italic>Ph. digitalis</italic> is a pathogen
of <italic>Digitalis</italic> spp. (<italic>Scrophulariaceae</italic>), but shares the feature
with <italic>Ph. bellidis</italic> that it is also recorded as a seed-pathogen
(<xref ref-type="bibr" rid="ref38">Boerema & Dorenbosch
1979</xref>). In contrast, <italic>Ph. senecionis</italic> is only known as a
necrophyte.</p><p><fig position="float" id="fig4"><label>Fig. 4.</label><caption><p><italic>Phoma pedeiae</italic> (<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=124517&link_type=cbs">CBS
124517</ext-link>). A–C. Fourteen-day-old colonies on OA (A), MEA (B)
and CHA (C). D. Pycnidia. E. Section of the pycnidial wall. F. Conidia. Scale
bars: D = 100 μm; E–F = 10 μm.</p></caption><graphic xlink:href="1fig4"></graphic></fig></p><p><italic><bold>Phoma bellidis</bold></italic> Neerg., Friesia 4: 74. 1950.</p><p><italic>Specimens examined</italic>: <bold>The Netherlands</bold>, from seed of
<italic>Bellis perennis</italic>, 1985, G.H. Boerema, CBS H-5200, culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=714.85&link_type=cbs">CBS 714.85</ext-link> = PD
74/265; from <italic>Bellis</italic> sp., 1994, J. de Gruyter, PD 94/886.</p><p><italic><bold>Phoma digitalis</bold></italic> Boerema apud Boerema & Dorenbosch,
Verslagen Meded. Plziektenk. Dienst Wageningen 153: 19. 1979.</p><p><italic>Specimen examined</italic>: <bold>The Netherlands</bold>, Ommen, from
<italic>Digitalis</italic> sp., 1990, J. de Gruyter,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=109179&link_type=cbs">CBS 109179</ext-link> = PD
90/835-1.</p><p><italic><bold>Phoma senecionis</bold></italic> P. Syd., Hedwigia, Beibl. 38: 136.
1899.</p><p><italic>Specimen examined</italic>: <bold>New Zealand</bold>, Raetihi, from a stem of
<italic>Senecio jacobaea</italic>, Feb. 1977, S. Ward,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=160.78&link_type=cbs">CBS 160.78</ext-link> = LEV
11451.</p></sec><sec><title>Group I:</title><p>Group I comprises three <italic>Phoma</italic> taxa (<italic>Ph. acetosellae, Ph.
macrostoma</italic> var. <italic>macrostoma</italic> and var. <italic>incolorata</italic>) that
were placed in the section <italic>Phyllostictoides</italic> on the basis of the
presence of septate conidia (<xref ref-type="bibr" rid="ref99">Gruyter
<italic>et al.</italic> 2002</xref>), but also accommodates <italic>Ph.
viburnicola.</italic> The placement of this species in section <italic>Phoma</italic> can
be debated, as a single septate conidium has been observed in strain
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=500.91&link_type=cbs">CBS 500.91</ext-link>, one of
the strains that was designated as reference strain
(<xref ref-type="bibr" rid="ref96">De Gruyter & Noordeloos
1992</xref>). Also <italic>D. exigua</italic>(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=183.55&link_type=cbs">CBS 183.55</ext-link> -
Neotype) is found in this clade, the type species of the genus
<italic>Didymella</italic> (<xref ref-type="bibr" rid="ref100">De Gruyter <italic>et
al.</italic> 2009</xref>), which does however not produce an anamorph state.
The four species do not exhibit a shared pathological feature or geographic
origin. The variety <italic>incolorata</italic> differs from var. <italic>macrostoma</italic>in lacking a red to violet pigment in the hyphae and any reaction to NaOH.</p><p><italic><bold>Phoma acetosellae</bold></italic> (A.L. Sm. & Ramsb.) Aa & Boerema,
in De Gruyter, Boerema & Van der Aa, Persoonia 18(1): 16. 2002.</p><p><italic>Basionym</italic>: <italic>Phyllosticta acetosellae</italic> A.L. Sm. & Ramsb.,
Trans. Brit. Mycol. Soc. 4: 173. 1912.</p><p><italic>Specimens examined</italic>: <bold>France</bold>, Corrèze, Monteil sur
Bois, from a leaf of <italic>Rumex acetosella</italic>, 1976, H.A. van der Aa, CBS
H-16138, culture 631.76. <bold>The Netherlands</bold>, Baarn, from a stem of
<italic>Rumex hydrolapathum</italic>, March 1996, H.A. van der Aa,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=179.97&link_type=cbs">CBS 179.97</ext-link>.</p><p><italic><bold>Phoma macrostoma</bold></italic> var. <italic><bold>incolorata</bold></italic> (A.S.
Horne) Boerema & Dorenb., Persoonia 6(1): 55. 1970.</p><p><italic>Basionym</italic>: <italic>Polyopeus purpureus</italic> var. <italic>incolorata</italic>A.S. Horne, J. Bot. 58: 240. 1920.</p><p><italic>Specimens examined</italic>: <bold>Switzerland</bold>, Vierwaldstättersee,
near Brunnen, from a leaf of <italic>Acer pseudoplatanus</italic>, Oct. 1968, J.
Gemmen, CBS H-20240, culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=223.69&link_type=cbs">CBS 223.69</ext-link>. <bold>The
Netherlands</bold>, from <italic>Malus sylvestris</italic>, 1983, J. de Gruyter,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=109173&link_type=cbs">CBS 109173</ext-link> = PD
83/908.</p><p><italic><bold>Phoma macrostoma</bold></italic> var. <italic><bold>macrostoma</bold></italic> Mont.,
Annls Sci. Nat., Bot. III 11: 52. 1849.</p><p><italic>Specimens examined</italic>: <bold>Germany</bold>, near München, from the
bark of <italic>Larix decidua</italic>, 1995, G.J. Verkley,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=482.95&link_type=cbs">CBS 482.95</ext-link>. <bold>The
Netherlands</bold>, Wageningen, from wood of <italic>Malus sylvestris</italic>, Sep.
1969, G.H. Boerema, CBS H-16431, culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=529.66&link_type=cbs">CBS 529.66</ext-link> = PD
66/521.</p><p><fig position="float" id="fig5"><label>Fig. 5.</label><caption><p><italic>Phoma dimorpha</italic> (<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=346.82&link_type=cbs">CBS
346.82</ext-link>). A–C. Fourteen-day-old colonies on OA (A), MEA (B)
and CHA (C). D. Pycnidia on stem of <italic>Urtica dioica</italic>. E. Pycnidia. F.
Pycnidial wall. G–H. Conidia <italic>in vitro</italic> (G) and <italic>in vivo</italic>(H). Scale bars: D–E = 100 μm; F = 20 μm; G–H = 10
μm.</p></caption><graphic xlink:href="1fig5"></graphic></fig></p><p><italic><bold>Phoma viburnicola</bold></italic> Oudem., Contr. Flora Mycol. d. Pays-Bas
17: 247. 1901.</p><p><italic>Specimens examined</italic>: <bold>The Netherlands</bold>, Wageningen, Aboretum,
from <italic>Viburnum cassioides</italic>, 1969, G.H. Boerema, CBS H-16605, culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=523.73&link_type=cbs">CBS 523.73</ext-link> = PD
69/800; from <italic>Chamaecyparis lawsoniana</italic>, 1981, G.H. Boerema,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=371.91&link_type=cbs">CBS 371.91</ext-link> = PD
81/413; Baarn, from a leaf of <italic>Ilex aquifolium</italic>, 1993, J. de Gruyter,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=500.91&link_type=cbs">CBS 500.91</ext-link> = PD
83/222.</p></sec><sec><title>Group J:</title><p>This small group (BPP = 1.00, RBS = 96 %) comprises only two species.
Because of the production of dictyochlamydospores, <italic>Phoma boeremae</italic> was
suggested to belong to the section <italic>Peyronellaea</italic> (Group K,
<xref ref-type="bibr" rid="ref19">Aveskamp <italic>et al</italic>.
2009a</xref>), to which the present group is closely related. No such
structures were, however, observed in its sister species, <italic>Ph.
dimorpha</italic> sp. nov. This species is known from a single strain, which
sporulates poorly and may be degenerated.</p><p><italic><bold>Phoma boeremae</bold></italic> Gruyter, Persoonia 18 (1): 91. 2002.</p><p><italic>Specimen examined</italic>: <bold>Australia</bold>, Victoria, Burnley Gardens,
from seed of <italic>Medicago littoralis</italic> cv. Harbinger, Febr. 1982, M.
Mebalds, <bold>neotype</bold> L 996.294.536, ex-neotype culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=109942&link_type=cbs">CBS 109942</ext-link> = PD
84/402.</p><p><italic><bold>Phoma dimorpha</bold></italic> Aveskamp, Gruyter & Verkley, <bold>sp.
nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515595&link_type=mb">MB515595</ext-link>.
<xref ref-type="fig" rid="fig5">Fig. 5</xref>.</p><p>Conidia dimorpha, in vitro cylindrica, glabra, hyalina, continua,
8–9.5(–10.5) × (2–)2.5–3(–3.5) μm,
(5–)6–8(–10) guttulis minutis apolaribus praedita, in vivo
eguttulata, (8–)9–12(–12.5) ×
(4.5–)5–5.5(–6.5) μm.</p><p><italic>Etymology</italic>: The epithet refers to the two different conidial types
that are observed.</p><p><italic>Pycnidia</italic> produced only scarcely <italic>in vitro</italic>, in clusters of
<italic>ca</italic>. 4–10 elements, globose, glabrous, non-papillate, produced
on the agar surface, relatively small, measuring
(65–)85–170(–190) μm diam. <italic>Ostioles</italic> single,
non-pappillate. <italic>Pycnidial wall</italic> pseudoparenchymatous, composed of
isodiametric cells, 4–7 layers, 14–20 μm thick.
<italic>Conidiogenous cells</italic> phialidic, hyaline, simple, smooth, flask-shaped,
<italic>ca.</italic> 5.5–7 × 4.5–6.5 μm. <italic>Conidia</italic>cylindrical, thin-walled, smooth, hyaline, aseptate 8–9.5(–10.5)
× (2–)2.5–3(–3.5) μm, with
(5–)6–8(–10) minute apolar guttules. <italic>In vivo</italic>eguttulate and somewhat broader, measuring (8–)9–12(–12.5)
× (4.5–)5–5.5(–6.5) μm. Conidial exudates not
observed.</p><p><italic>Culture characteristics</italic>: Colonies on OA 45–50 mm diam after
7 d, margin regular. Immersed mycelium olivaceous-black, in some sectors
covered by a low mat of floccose white to grey aerial mycelium, towards colony
margin the aerial mycelium is gradually becoming more felted and white;
reverse olivaceous buff to dark mouse-grey. Colonies on MEA 50–55 mm
diam after 7 d, margin regular. Immersed mycelium hyaline, amber or iron-grey.
Only sparsely small white tufts of whitish aerial mycelium are produced in
older cultures; reverse concolourous. Colonies on CHA 55–60 mm diam
after 7 d, margin regular. Immersed mycelium hyaline, honey to isabelline or
dark mouse-grey. Aerial mycelium more proliferent near colony margin initially
white, later developing to iron-grey with olivaceous-grey tinges; reverse
black, but hyaline near colony centre. Application of NaOH did not have any
effect. In older cultures white dendritic crystals are formed both in the
aerial mycelium and in immersed in the agar.</p><p><italic>Specimen examined</italic>: <bold>Spain</bold>, Canary Isles, Gran Canaria, from
phyllocladium of <italic>Opuntiae</italic> sp., June 1982, H.A. Van der Aa,
<bold>holotype designated here</bold> CBS H-20234, culture ex-holotype
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=346.82&link_type=cbs">CBS 346.82</ext-link>.</p><p><italic>Notes</italic>: Although sufficient pycnidial primordia are formed on OA,
maturation of pycnidia is only incidentally observed <italic>in vitro</italic>.
Therefore the characters of the pycnidia and the pycnidial wall described here
are based on only three samples. Formation of mature pycnidia can be induced
by addition of a sterilised stem piece of stinging nettle (<italic>Urtica
dioica</italic>). The conidia that were described from <italic>in vivo</italic> material
were obtained using this technique.</p><p>Several other <italic>Phoma</italic> species are known from <italic>Opuntiae</italic>,
including <italic>Ph. opuntiae</italic> (<italic>Phoma sensu lato</italic>) and <italic>Ph.
longicolla</italic> sp. nov. (see below). The conidia of <italic>Ph. opuntiae</italic>are, however, considerably smaller, measuring 2.5–3.5 ×
1–1.5 μm (<xref ref-type="bibr" rid="ref96">De Gruyter &
Noordeloos 1992</xref>), whereas the main difference with <italic>Ph.
longicolla</italic> are the pycnidia, which are uniostiolate and significantly
larger in the latter species.</p></sec><sec><title>Group K – Peyronellaea:</title><p>This group (BPP = 1.00, RBS = 58 %) comprises many of the chlamydospore
forming species, including the majority of the species that were accommodated
in <italic>Phoma</italic> section <italic>Peyronellaea</italic> (Boerema <italic>et al.</italic><xref ref-type="bibr" rid="ref43">1965a</xref>,
<xref ref-type="bibr" rid="ref45">1968</xref>,
<xref ref-type="bibr" rid="ref46">1971</xref>,
<xref ref-type="bibr" rid="ref47">1973</xref>,
<xref ref-type="bibr" rid="ref48">1977</xref>). Also <italic>Ph.
glomerata</italic>, type species of this section is accommodated here
(<xref ref-type="bibr" rid="ref34">Boerema 1997</xref>). However, as
section, <italic>Peyronellaea</italic> has a polyphyletic nature
(<xref ref-type="bibr" rid="ref19">Aveskamp <italic>et al.</italic>2009a</xref>). <italic>Phoma chrysanthemicola, Ph. violicola</italic> and the
recently established species <italic>Ph. schachtii</italic>(<xref ref-type="bibr" rid="ref19">Aveskamp <italic>et al.</italic>2009a</xref>) have been found to be basal to the <italic>Didymellaceae</italic>(<xref ref-type="fig" rid="fig1">Fig. 1</xref>), whilst several
species producing botryoid chlamydospores, representing the genus
<italic>Epicoccum</italic> as emended below, are clustered in group M. Also <italic>Ph.
infossa</italic> and <italic>Ph. omnivirens</italic>, which have proven to produce
dictyochlamydospores in culture (<xref ref-type="bibr" rid="ref19">Aveskamp
<italic>et al.</italic> 2009a</xref>), are not situated in this part of the
phylogenetic tree. <italic>Peyronellaea calidophila</italic> and <italic>Ph.
microchlamydospora</italic> reside in the basal lineages of this clade.</p><p>Also several <italic>Phoma</italic> species that were not included in
<italic>Peyronellaea</italic> in the Boeremaean taxonomical system, but that do
produce either uni- or multicellular chlamydospores, are included in this
clade. In <italic>Ph. gardeniae, Ph. narcissi</italic>, and <italic>Ph. zeae-maydis</italic>multicellular chlamydospores have been observed, whereas <italic>Ph. pinodella,
Ph. arachidicola</italic>, and <italic>Ph. heteroderae</italic> are species that form
unicellular chlamydospores. Several species in this clade, however, have never
been recorded to produce any unicellular or multicellular chlamydospores.
These species are <italic>Ph. alectorolophi, Ph. obtusa</italic> and <italic>Ph.
protuberans</italic>, which will be treated in a subsequent section of this
paragraph, and <italic>Ph. anserina, Ph. aurea, Ph. nigricans</italic> and <italic>Ph.
eucalyptica</italic>. However, two of these species, <italic>Ph. anserina</italic> and
<italic>Ph. eucalyptica</italic> are well-known for the formation of swollen cells and
anastomosis in culture (<xref ref-type="bibr" rid="ref96">De Gruyter &
Noordeloos 1992</xref>), which may be regarded as a precursor to
chlamydospore formation. The ancestral location of <italic>Ph. anserina</italic> in
this clade may also be an indication that chlamydospore production has not
completely been developed yet in this group. The high posterior probabillity
for this group justifies the recognition of a separate genus in the
<italic>Didymellaceae</italic>. Therefore the genus name <italic>Peyronellaea</italic> Goid.
is re-established, and the associated species are recombined into this genus
below.</p><p>The plurivorous species <italic>Ph. calorpreferens</italic> and <italic>Ph.
heteroderae</italic> share identical LSU, ITS and TUB genes. Also morphologically
the representative strains of these species are highly similar. A
synonymisation of these species is therefore proposed in this paper.</p><p>Another notable subgroup within this clade is a cluster formed by
<italic>Didymella pinodes, D. lethalis, D. arachidicola</italic> and <italic>Ph.
pinodella</italic>. Recently, Irinyi <italic>et al.</italic>(<xref ref-type="bibr" rid="ref117">2009</xref>) synonymised <italic>Ph.
sojicola</italic> with <italic>Ph. pinodella</italic>, based on morphological observations
and sequence data of ITS, β-tubulin and translation elongation factor
1-α. This indicates that the notorious pathogen of green pea (<italic>Pisum
sativum</italic>) is also capable of infecting soybean (<italic>Glycine max</italic>).
These observations are supported by the results obtained in the present study.
As reported in previous studies
(<xref ref-type="bibr" rid="ref87">Faris-Mokaiesh <italic>et al.</italic>1996</xref>, <xref ref-type="bibr" rid="ref155">Onfroy <italic>et al.</italic>1999</xref>, <xref ref-type="bibr" rid="ref88">Fatehi <italic>et al.</italic>2003</xref>, <xref ref-type="bibr" rid="ref163">Peever <italic>et al.</italic>2007</xref>), <italic>Ph. pinodella</italic> appears to be very closely related
to <italic>D. pinodes</italic> (anam. <italic>Ascochyta pinodes</italic>) and because these
species share the same host range they are often confused. Both species can
however easily be differentiated on basis of the amount of septate conidia
formed <italic>in vitro</italic>, abundantly in <italic>D. pinodes</italic>, and in very small
numbers in <italic>Ph. pinodella</italic>.</p><p>Because <italic>Ph. pinodella</italic> is morphologically so similar to <italic>Ph.
medicaginis</italic>, it was once regarded as a variety of this species by Boerema
<italic>et al.</italic> (<xref ref-type="bibr" rid="ref44">1965b</xref>). The
variety was elevated to species rank after careful observation
(<xref ref-type="bibr" rid="ref233">White & Morgan-Jones
1987</xref>), but the varietal name is however currently still in common
use (<italic>e.g.</italic> <xref ref-type="bibr" rid="ref155">Onfroy <italic>et al</italic>.
1999</xref>, <xref ref-type="bibr" rid="ref88">Fatehi <italic>et al</italic>.
2003</xref>, <xref ref-type="bibr" rid="ref223">Taylor & Ford
2007</xref>). The results obtained in this study however, illustrate a
substantial phylogenetical distance to <italic>Ph. medicaginis</italic>, and warrant
recognition at species level, in the in the re-instated genus
<italic>Peyronellaea</italic>.</p><p>The close association of <italic>Ph. arachidicola</italic> with <italic>Ph.
pinodella</italic> and <italic>D. pinodes</italic> is reflected by the morphology of these
species, which all produce, next to septate and aseptate conidia, also globose
to ellipsoidal unicellular chlamydospores, which may be formed in chains.
These chlamydospores measure 5–20 μm diam, which is somewhat larger
than the species in group N. The close relationship of these three species has
been hypothesised before, and was based on chemical analysis of the crystals
produced by these taxa (<xref ref-type="bibr" rid="ref152">Noordeloos <italic>et
al.</italic> 1993</xref>).</p><p><italic>Didymella arachidicola</italic> is a specific pathogen of groundnut
(<italic>Arachis hypogaea</italic>), another host plant of the family
<italic>Fabaceae</italic> with which the other species in this subclade are also
associated.</p><p>In the <italic>Ph. pinodella</italic> / <italic>D. pinodes</italic> subcluster (BPP = 1.00,
RBS = 94 %), four teleomorph species are found with a coelomycete anamorph
state. Next to <italic>D. pinodes</italic>, these are <italic>D. alectorolophi, D.
arachidicola,</italic> and <italic>D. lethalis</italic>. A fifth teleomorph is the sexual
state of <italic>Ph. pinodella</italic> (as <italic>Ph. medicaginis</italic> var.
<italic>pinodella</italic>) that is reported and described by Bowen <italic>et al.</italic>(<xref ref-type="bibr" rid="ref56">1997</xref>), but that has not been
named thus far. From a phylogenetic point of view, this record is very
plausible as all species in the subclade in which <italic>Ph. pinodella</italic> is
embedded, do form a <italic>Didymella</italic>-like teleomorph
(<xref ref-type="fig" rid="fig2">Fig. 2</xref>). However, as we did
not include mating type tests in our studies, and as the species is probably
heterothallic (<xref ref-type="bibr" rid="ref56">Bowen <italic>et al.</italic>1997</xref>), pseudothecia were not observed in the present study. A
formal name for the teleomorph of <italic>Ph. pinodella</italic> could therefore not
be proposed here either.</p><p>A fifth species in group K that has a known teleomorph is <italic>Ph.
zeae-maydis</italic>. This species is however only distantly related to the four
species mentioned above. Nevertheless, it can be concluded that the sole
teleomorph genus that is associated with group K is <italic>Didymella</italic>-like.
This would further support the suggestion
(<xref ref-type="bibr" rid="ref163">Peever <italic>et al.</italic> 2007</xref>)
that the teleomorph name for <italic>A. pinodes</italic> that is often referred to by
plant pathologists, <italic>Mycosphaerella pinodes,</italic> should be omitted.</p><p>Remarkably, three species are found in this clade that are identical based
on sequence analyses, but that are morphologically rather distinct. Also
sequence comparisons of parts of the actin and calmodulin genes did not reveal
any differences between those four strains (Aveskamp & Woudenberg, unpubl.
data). <italic>Phoma alectorolophi</italic> and <italic>Ph. protuberans</italic> are
associated with <italic>Phoma</italic> section <italic>Sclerophomella</italic>(<xref ref-type="bibr" rid="ref54">Boerema <italic>et al.</italic> 1997</xref>,
<xref ref-type="bibr" rid="ref99">De Gruyter <italic>et al.</italic>2002</xref>), because of the thick-walled pycnidia formed in culture and
<italic>in vivo</italic>. However, because of the production of relatively large
secondary conidia, a link with sections <italic>Heterospora</italic> or
<italic>Phyllostictoides</italic> can also be advocated. Colony characters,
microscopic features and ecology indicate that the two species should actually
be rather distinct. A third taxon found in this group is <italic>Ph. obtusa</italic>,
a saprobic species that has a thin pycnidial wall and lacks septate conidia.
Nevertheless, these three species are recovered in a clade in which solely
chlamydospore-forming species reside, a character that never has been recorded
in any of these taxa. The explanation of the contrast between the level of
genetic and morphological similarity will be one of the main challenges in
<italic>Phoma</italic> taxonomy.</p><p><italic><bold>Peyronellaea</bold></italic> Goid. ex Togliani, Ann. Sperim. Agrar. II 6:
93. 1952, <bold>emend</bold><italic>.</italic> Aveskamp, Gruyter & Verkley</p><p><italic>Conidiomata</italic> pycnidial, globose to subglobose, measuring
50–380 μm diam, on agar surface or immersed, solitary or confluent,
ostiolate or poroid. <italic>Pycnidial wall</italic> pseudoparenchymatous, counting
2–8 cell layers of which the outer 1–3 are brown or olivaceous
pigmented. <italic>Conidiogenous cells</italic> phialidic, hyaline, simple, smooth,
ampulliform or doliiform, <italic>ca</italic>. 3.5–7 × 3.5–6 μm.
<italic>Conidia</italic> generally aseptate, ellipsoidal to subglobose, thin-walled,
smooth, hyaline, but in older cultures conidia may become pigmented, generally
measuring 4–15 × 2–4 μm, but larger or septated conidia
may occur in at least one species. <italic>Unicellular chlamydospores</italic> often
abundantly formed in and on the agar and in the aerial mycelium, globose,
intercalary, brown or olivaceous pigmented, measuring 5–22 μm diam.
<italic>Multicellular chlamydospores</italic> mainly alternarioid, terminal or
intercalary, often in chains, brown or olivaceous pigmented, 10–50
× 7–25 μm. <italic>Pseudothecia</italic> only present in a few species,
(sub-)globose, up to 200 μm diam, but in one species also flattened
pseudothecia occur. <italic>Asci</italic> cylindrical to clavate, measuring
35–65 × 11–17 μm, always 8-spored, biseriate.
<italic>Ascospores</italic> ellipsoid, measuring 12–24 × 4–8 μm,
uniseptate, upper cell usually larger than the lower cells.</p><p><italic>Type species</italic>: <italic>Peyronellaea glomerata</italic> (Corda) Goid. ex
Togliani</p><p><italic><bold>Peyronellaea alectorolophi</bold></italic> (Rehm.) Aveskamp, Gruyter &
Verkley, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515597&link_type=mb">MB515597</ext-link>.</p><p><italic>Basionym</italic>: <italic>Didymella alectorolophi</italic> Rehm, apud Ade,
Hedwigia 64: 294. 1923.</p><p><list list-type="simple"><list-item><p>≡ <italic>Phoma alecotorolophi</italic> Boerema, Gruyter & Noordel.,
Persoonia 16(3): 366. 1997.</p></list-item></list></p><p><italic>Specimen examined</italic>: <bold>The Netherlands</bold>, from seed of
<italic>Rhinanthus major</italic>, 1993, L 992.167.515, culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=132.96&link_type=cbs">CBS 132.96</ext-link> = PD
93/853.</p><p><italic><bold>Peyronellaea americana</bold></italic> (Morgan-Jones & J.F. White)
Aveskamp, Gruyter & Verkley, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515596&link_type=mb">MB515596</ext-link>.</p><p><italic>Basionym</italic>: <italic>Phoma americana</italic> Morgan-Jones & J.F. White,
Mycotaxon 16(2): 406. 1983.</p><p><italic>Specimens examined</italic>: <bold>Argentina</bold>, Buenos Aires Province,
Olavarria, from leaves of <italic>Triticum aestivum</italic> cv. Buck Diamante, Aug.
2002, A. Perelló, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=112525&link_type=cbs">CBS
112525</ext-link>. <bold>Denmark</bold>, Copenhagen, from seed of <italic>Phaseolus
vulgaris</italic>, May 1965, S.B. Mathur,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=256.65&link_type=cbs">CBS 256.65</ext-link>.
<bold>Nigeria</bold>, from <italic>Sorghum vulgare</italic>, 1979, PD 79/58. <bold>South
Africa</bold>, from <italic>Zea mays</italic>, 1978, PD 78/1059. <bold>U.S.A.</bold>, Arkansas,
from pod lesions of <italic>Glycine max</italic>, 1981, H.J. Walters,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=568.9797&link_type=cbs">CBS 568.9797</ext-link> =
ATCC 44494 = PD 94/1544; Georgia, from <italic>Zea mays</italic>, 1985, G.H. Boerema,
CBS H-16144, culture <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=185.85&link_type=cbs">CBS
185.85</ext-link> = PD 80/1191; from <italic>Zea mays</italic>, 1980, PD 80/1143.
Unknown origin, from a nematode cyst, 1982, G.H. Boerema, PD 82/1059.</p><p><italic><bold>Peyronellaea anserina</bold></italic> <bold>(</bold>Marchal) Aveskamp, Gruyter
& Verkley, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515598&link_type=mb">MB515598</ext-link>.</p><p><italic>Basionym</italic>: <italic>Phoma anserina</italic> Marchal, Champignon Copr. 11:
1891.</p><p><italic>Specimens examined</italic>: <bold>The Netherlands</bold>, from <italic>Pisum
sativum</italic>, 1979, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=363.91&link_type=cbs">CBS
363.91</ext-link> = PD 79/712; Ter Apel, from potato flour, 1983,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=360.84&link_type=cbs">CBS 360.84</ext-link>.</p><p><italic><bold>Peyronellaea arachidicola</bold></italic> (Khokhr.) Aveskamp, Gruyter &
Verkley, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515599&link_type=mb">MB515599</ext-link>.</p><p><italic>Basionym</italic>: <italic>Mycosphaerella arachidicola</italic> Khokhr., Bolezni i
vrediteli maslichnykh kul'tur 1(2): 29. 1934.</p><p><list list-type="simple"><list-item><p>≡ <italic>Didymella arachidicola</italic> (Khokhr.) Tomilin, Opredelitel'
gribov roda Mycosphaerella Johans: 285. 1979.</p></list-item></list></p><p><italic>Anamorph</italic>: <italic>Phoma arachidicola</italic> Marasas, Pauer &
Boerema, Phytophylactica 6(3): 200. 1974.</p><p><italic>Specimens examined</italic>: <bold>South Africa</bold>, Cape Province, Jan
Kempdorp, Vaalharts Research Station, from a leaf of <italic>Arachis
hypogaea</italic>, Mar. 1972, G.D. Pauer, <bold>isotype</bold> of <italic>Ph.
arachidicola</italic> CBS H-7601, ex-isotype culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=333.75&link_type=cbs">CBS 333.75</ext-link> = ATCC
28333 = IMI 386092 = PREM 44889; <bold>Zimbabwe</bold>, from <italic>Arachis
hypogaea</italic>, 1980, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=315.90&link_type=cbs">CBS
315.90</ext-link> = PD 80/1190.</p><p><italic><bold>Peyronellaea aurea</bold></italic> (Gruyter, Noordel. & Boerema)
Aveskamp, Gruyter & Verkley, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515600&link_type=mb">MB515600</ext-link>.</p><p><italic>Basionym</italic>: <italic>Phoma aurea</italic> Gruyter, Noordel. & Boerema,
Persoonia 15(3): 394. 1993.</p><p><italic>Specimen examined</italic>: <bold>New Zealand</bold>, Auckland, from a stem of
<italic>Medicago polymorpha</italic>, 1978, <bold>holotype</bold> L 992.177.422, ex-holotype
culture <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=269.93&link_type=cbs">CBS 269.93</ext-link>= PD 78/1087.</p><p><italic><bold>Peyronellaea australis</bold></italic> Aveskamp, Gruyter & Verkley,
<bold>nom. nov.</bold> pro <italic>Phoma nigricans</italic> P.R. Johnst. & Boerema,
MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515601&link_type=mb">MB515601</ext-link>.</p><p><list list-type="simple"><list-item><p>≡ <italic>Phoma nigricans</italic> P.R. Johnst. & Boerema, New Zealand J.
Bot. 19(4): 394. 1982.</p></list-item></list></p><p><italic>Etymology</italic>: Epithet refers to the Southern Hemisphere, where this
fungus is mainly found.</p><p><italic>Specimens examined</italic>: <bold>New Zealand</bold>, from <italic>Actinidea
chinensis</italic>, 1977, P.R. Johnston, PD 77/919; Auckland, Mt. Albert, from a
leaf of <italic>Actinidia chinensis</italic>, Apr. 1979, P.R. Johnston, <bold>isotype</bold>CBS H-7619, ex-isotype culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=444.81&link_type=cbs">CBS 444.81</ext-link> = PDDCC
6546.</p><p><italic>Note</italic>: A new name was sought for this species, as the epithet
“<italic>nigricans</italic>” already was occupied in
<italic>Peyronellaea</italic>, referring to a species which is now synonymised with
<italic>Pey. pomorum</italic> var. <italic>circinata</italic> (see below).</p><p><italic><bold>Peyronellaea calorpreferens</bold></italic> (Boerema, Gruyter &
Noordel.) Aveskamp, Gruyter & Verkley, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515602&link_type=mb">MB515602</ext-link>.</p><p><italic>Basionym</italic>: <italic>Phoma pomorum</italic> var. <italic>calorpreferens</italic>Boerema, Gruyter & Noordel. apud Boerema, Persoonia 15: 207. 1993.</p><p><list list-type="simple"><list-item><p>≡ <italic>Phoma calorpreferens</italic> (Boerema, Gruyter & Noordel.)
Aveskamp, Gruyter & Verkley, Mycologia 101: 370. 2009.</p></list-item><list-item><p>= <italic>Phoma heteroderae</italic> Sen Y. Chen, D.W. Dicks. & Kimbr.,
Mycologia 88: 885.1996.</p></list-item></list></p><p><italic>Conidiomata</italic> pycnidial, solitary or confluent, partially or
completely immersed in the agar, (sub-)globose or irregular due to the
presence of 1(–4) slightly papillate ostioles, measuring
(70–)100–200(–250) μm diam. <italic>Pycnidial wall</italic>pseudoparenchymatous, composed of isodiametric cells, 2–5 layers thick,
with many hyphal outgrows, some setae-like. <italic>Conidiogenous cells</italic>phialidic, hyaline, simple, smooth, flask-shaped, <italic>ca</italic>. 3–5.5
× 3–6.5 μm. <italic>Conidia</italic> broadly ellipsoidal to ovoid to
cylindrical, thin-walled, smooth, hyaline, (3.5–)4–8.5(–12)
× 2–3.5(–4.5) μm, aseptate, with
(1–)2–5(–8) polar guttules. Conidial matrix pale pink.
<italic>Chlamydospores</italic> highly variable in shape and size, mostly unicellular
but also multicellular. Where unicellular, pale brown to brown, guttulate,
intercalary, solitary or in chains, globose, 7.5–19(–26) μm,
thick-walled and often with a distinct “envelope”. Where
multicellular dictyosporous alternarioid or botryoid, brown to black, terminal
or occasionally intercalary in chains of unicellular chlamydospores, measuring
<italic>ca.</italic> (16–)21–55 × (7–)12–30(–33)
μm.</p><p><italic>Specimens examined</italic>: <bold>The Netherlands</bold>, from undefined food
material, 1973, G.H. Boerema, <bold>holotype</bold> L 990.290.418, ex-holotype
culture <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=109.92&link_type=cbs">CBS 109.92</ext-link>= PD 73/1405. <bold>U.S.A.</bold>, Florida, Gainesville, from eggs of <italic>Heterodera
glycines</italic> from greenhouse soil,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=630.97&link_type=cbs">CBS 630.97</ext-link> = ATCC
96683 = IMI 361196 = PD 96/2022; from indoor environment, 1993,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=875.97&link_type=cbs">CBS 875.97</ext-link> = PD
93/1503.</p><p><italic>Notes</italic>: <italic>Peyronellaea calorpreferens</italic> is a taxon that was
recently elevated from variety level to species rank, as <italic>Phoma
calorpreferens</italic> (<xref ref-type="bibr" rid="ref19">Aveskamp <italic>et
al.</italic> 2009a</xref>). Due to its morphological and genetical similarity
with <italic>Ph. heteroderae</italic>, it is concluded that both taxa are actually one
and the same species. According to the International code of Botanical
Nomenclature (McNeal <italic>et al.</italic> 2006) the epithet <italic>calorpreferens</italic>has priority, as its basionym <italic>Ph. pomorum</italic> var.
<italic>calorpreferens</italic> was published earlier.</p><p>The type of <italic>Peyronellaea calorpreferens</italic> has been recovered from
food materials, but Boerema
(<xref ref-type="bibr" rid="ref33">1993</xref>) hypothesises about the
plurivorous nature of this taxon, and mainly records it as a worldwide
occurring soil- and seedborne opportunist, whereas Chen <italic>et al.</italic>(<xref ref-type="bibr" rid="ref64">1996</xref>) record this species (as
<italic>Ph. heteroderae</italic>) from eggs of a cyst nematode, <italic>Heterodera
glycines</italic>.</p><p><italic><bold>Peyronellaea coffeae-arabicae</bold></italic> (Aveskamp, Verkley &
Gruyter) Aveskamp, Gruyter & Verkley, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515603&link_type=mb">MB515603</ext-link>.</p><p><italic>Basionym</italic>: <italic>Phoma coffeae-arabicae</italic> Aveskamp, Verkley &
Gruyter, Mycologia 101(3): 371. 2009.</p><p><italic>Specimens examined</italic>: <bold>Ethiopia</bold>, from <italic>Coffea arabica</italic>,
1984, M.M.J. Dorenbosch, <bold>holotype</bold> CBS H-20143, ex-holotype culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123380&link_type=cbs">CBS 123380</ext-link> = PD
84/1013; from <italic>Coffea arabica</italic>, 1984, M.M.J. Dorenbosch,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123398&link_type=cbs">CBS 123398</ext-link> = PD
84/1014.</p><p><italic><bold>Peyronellaea curtisii</bold></italic> (Berk.) Aveskamp, Gruyter &
Verkley, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515604&link_type=mb">MB515604</ext-link>.</p><p><italic>Basionym</italic>: <italic>Hendersonia curtisii</italic> Berk., in Cooke, Nuovo
Giorn. Bot. Ital. 10: 19. 1878.</p><p><list list-type="simple"><list-item><p>≡ <italic>Stagonosporopsis curtisii</italic> (Berk.) Boerema, in Boerema
& Dorenbosch, Verslagen Meded. Plziektenk. Dienst Wageningen 157: 20.
1981.</p></list-item><list-item><p>= <italic>Phyllosticta narcissi</italic> Aderh., Centralbl. Bakteriol., 2 Abth. 6:
632. 1900.</p></list-item><list-item><p>≡ <italic>Phoma narcissi</italic> (Aderh.) Boerema, Gruyter & Noordel.,
Persoonia 15(2): 215. 1993.</p></list-item></list></p><p><italic>Specimens examined</italic>: <bold>The Netherlands</bold>, from <italic>Nerine</italic>sp., May 1992, J. de Gruyter, culture 251.92 = PD 86/1145; from
<italic>Sprekelia</italic> sp., PD 92/1460. Unknown origin, from <italic>Ismene</italic> sp.,
1971, PD 71/6. Unknown origin, from <italic>Hippeastrum</italic> sp., 1976, PD
76/61.</p><p><italic><bold>Peyronellaea eucalyptica</bold></italic> (Sacc.) Aveskamp, Gruyter &
Verkley, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515605&link_type=mb">MB515605</ext-link>.</p><p><italic>Basionym</italic>: <italic>Phoma eucalyptica</italic> Sacc., Syll. Fung. 3: 78.
1884.</p><p><italic>Specimens examined</italic>: <bold>Australia</bold>, Western Australia, from a
leaf of <italic>Eucalyptus</italic> sp., 1979,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=377.91&link_type=cbs">CBS 377.91</ext-link> = PD
79/210. <bold>Croatia</bold>, Adriatic Sea, from seawater, 1973,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=508.91&link_type=cbs">CBS 508.91</ext-link> = PD
73/1413. <bold>Indonesia</bold>, Sumatra, Sulavesi, from <italic>Eugenia aromatica,</italic>1982, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=378.91&link_type=cbs">CBS 378.91</ext-link> =
PD 82/107.</p><p><italic><bold>Peyronellaea gardeniae</bold></italic> (S. Chandra & Tandon) Aveskamp,
Gruyter & Verkley, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515606&link_type=mb">MB515606</ext-link>.</p><p><italic>Basionym</italic>: <italic>Pyrenochaeta gardeniae</italic> S. Chandra & Tandon,
Mycopathol. Mycol. Appl. 29: 274. 1966.</p><p><list list-type="simple"><list-item><p>≡ <italic>Phoma gardeniae</italic> (S. Chandra & Tandon) Boerema, in
Boerema & Dorenbosch, Verslagen Meded. Plziektenk. Dienst Wageningen 156:
27. 1980.</p></list-item></list></p><p><italic>Specimens examined</italic>: <bold>India</bold>, Allahabad, from the leaf of
<italic>Gardenia jasminoides</italic>, 1966, S. Chandra and R.N. Tandon,
<bold>isotype</bold> CBS H-7605, ex-isotype culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=626.68&link_type=cbs">CBS 626.68</ext-link> = IMI
108771. <bold>Netherlands Antilles</bold>, Curacao, from air sample, 1978, A.
Kikstra, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=302.79&link_type=cbs">CBS 302.79</ext-link>= PD 79/1156.</p><p><italic><bold>Peyronellaea glomerata</bold></italic> (Corda) Goid. ex Togliani, Ann.
Sperim. Agrar. III 6: 93. 1952.</p><p><italic>Basionym</italic>: <italic>Coniothyrium glomeratum</italic> Corda, Icon. Fung.
(Prague) 4: 39. 1840.</p><p><list list-type="simple"><list-item><p>≡ <italic>Phoma glomerata</italic> (Corda) Wollenw. & Hochapfel, Z.
Parasitenk. 3(5): 592. 1936.</p></list-item></list></p><p><italic>Specimens examined</italic>: <bold>Germany</bold>, Berlin-Zehlendorf, Domäne
Düppel, from a tuber of <italic>Solanum tuberosum</italic>, 1936, H.W.
Wollenweber, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=293.36&link_type=cbs">CBS
293.36</ext-link> = MUCL 9882; Monheim, from <italic>Hordeum sativum</italic>, 1984,
M. Hossfeld, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=834.84&link_type=cbs">CBS
834.84</ext-link>; from indoor environment, 2003, C. Rudolph,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=112448&link_type=cbs">CBS 112448</ext-link>.
<bold>Romania</bold>, Bukarest, from a church wall-fresco, Nov. 1971, I. Ionita,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=133.96&link_type=cbs">CBS 133.96</ext-link> = PD
79/127. <bold>Russia</bold>, Novosibirsk, Hortus Botanicus, from a leaf of
<italic>Populus nigra</italic>, 1963, T.T. Kuznetsova,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=284.76&link_type=cbs">CBS 284.76</ext-link> = ATCC
26238 = IMI 176748 = VKM F-1842; Novosibirsk, Hortus Botanicus, from a leaf of
<italic>Rubus idaeus</italic>, 1963, T.T. Kuznetsova,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=287.76&link_type=cbs">CBS 287.76</ext-link> = ATCC
26240 = IMI 176746 = VKM F-1847; Novosibirsk, Hortus Botanicus, from a leaf of
<italic>Populus alba</italic>, 1963, T.T. Kuznetsova,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=288.76&link_type=cbs">CBS 288.76</ext-link> = ATCC
26243 = VKM F-1845; Novosibirsk, Hortus Botanicus, from a leaf of <italic>Allium
nutans</italic>, 1963, T.T. Kuznetsova,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=289&link_type=cbs">CBS 289</ext-link>. 76 = ATCC
26239 = IMI 176745 = VKM F-1846; Novosibirsk, Hortus Botanicus, from a leaf of
<italic>Ribes nigrum</italic>, 1963, T.T. Kuznetsova,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=290.76&link_type=cbs">CBS 290.76</ext-link> = ATCC
26244 = IMI 176747 = VKM F-1848; from <italic>Heracleum</italic> sp., 1973, PD
73/1415. <bold>The Netherlands</bold>, from a root of <italic>Lycopersicon
esculentum</italic>, 1949, D. Verleur,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=304.49&link_type=cbs">CBS 304.49</ext-link> = MUCL
9884; from <italic>Chrysanthemum</italic> sp., 1963,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=528.66&link_type=cbs">CBS 528.66</ext-link> = PD
63/590; from indoor bathroom environment, 1997, M. Komen,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=464.97&link_type=cbs">CBS 464.97</ext-link>; from
<italic>Medicago sativa</italic>, PD 77/47. <bold>U.K.</bold>, from air, PD 74/1023.
<bold>U.S.A.</bold>, Virginia, from <italic>Juniperus</italic> sp., Jan. 2002, A.Y, Rossman,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=120109&link_type=cbs">CBS 120109</ext-link>. Unknown
origin, from <italic>Cucumis sativus</italic>, PD 81/767; from <italic>Capsicum</italic> sp.,
PD 83/782.</p><p><italic><bold>Peyronellaea lethalis</bold></italic> (Ellis & Bartholomew) Aveskamp,
Gruyter & Verkley, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515607&link_type=mb">MB515607</ext-link>.</p><p><italic>Basionym</italic>: <italic>Ascochyta lethalis</italic> Ellis & Bartholomew,
Fungi Columb. 1808. 1903.</p><p><list list-type="simple"><list-item><p>= <italic>Mycosphaerella lethalis</italic> R. Stone, Ann. Mycol. 10: 587. 1912.</p></list-item><list-item><p>≡ <italic>Didymella lethalis</italic> (R. Stone) Sivan., Bitunicate
Ascomycetes and their Anamorphs: 424. 1984.</p></list-item></list></p><p><italic>Specimen examined</italic>: Unknown origin and substrate, 1925, A.W.
Archer, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=103.25&link_type=cbs">CBS
103.25</ext-link>.</p><p><italic><bold>Peyronellaea musae</bold></italic> P. Joly, <italic>Revue Mycol</italic>.
<bold>26</bold>: 97. 1961.</p><p><list list-type="simple"><list-item><p>≡ <italic>Phoma jolyana</italic> Piroz. & Morgan-Jones, Trans. Brit.
Mycol. Soc. 51: 200. 1968.</p></list-item></list></p><p><italic>Specimens examined</italic>: <bold>India</bold>, from fruit of <italic>Mangifera
indica</italic>, May 1969, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=463.69&link_type=cbs">CBS
463.69</ext-link>; from <italic>Malus sylvestris</italic>, PD 83/326.</p><p><italic>Notes</italic>: <italic>Phoma jolyana</italic> was originally described in the
genus <italic>Peyronellaea</italic>, as <italic>Pey. musae</italic>. The epithet
“<italic>jolyana</italic>” was later proposed for this species, as the
epithet <italic>musae</italic> was already occupied in <italic>Phoma</italic>(<xref ref-type="bibr" rid="ref164">Pirozynski & Morgan-Jones
1968</xref>). Here, we reinstate this fungus under its original name.</p><p><italic><bold>Peyronellaea obtusa</bold></italic> (Fuckel) Aveskamp, Gruyter &
Verkley, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515608&link_type=mb">MB515608</ext-link>.</p><p><italic>Basionym</italic>: <italic>Phoma obtusa</italic> Fuckel, Jahrb. Nassauischen
Vereins Naturk. 23–24: 378. 1870.</p><p><italic>Specimens examined</italic>: <bold>The Netherlands</bold>, from a root of
<italic>Daucus carota</italic>, July 1993, J. de Gruyter,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=377.93&link_type=cbs">CBS 377.93</ext-link> = PD
80/976; from <italic>Spinacia oleracea</italic>, July 1993, J. de Gruyter,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=391.93&link_type=cbs">CBS 391.93</ext-link> = PD
80/87.</p><p><italic><bold>Peyronellaea pinodella</bold></italic> (L.K. Jones) Aveskamp, Gruyter &
Verkley, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515609&link_type=mb">MB515609</ext-link>.</p><p><italic>Basionym</italic>: <italic>Ascochyta pinodella</italic> L.K. Jones, Bull. New York
State Agric. Exp. Sta. 547: 10. 1927.</p><p><list list-type="simple"><list-item><p>≡ <italic>Phoma medicaginis</italic> var. <italic>pinodella</italic> (L. K. Jones)
Boerema apud Boerema, Dorenbosch & Leffring, Netherlands J. Pl. Pathol.
71<bold>:</bold> 88. 1965.</p></list-item><list-item><p>≡ <italic>Phoma pinodella</italic> <bold>(</bold>L.K. Jones) Morgan-Jones &
K.B. Burch, Mycotaxon 29: 485. 1987.</p></list-item></list></p><p><italic>Specimens examined</italic>: <bold>Hungary</bold>, from <italic>Glycine max,</italic>1996, G. Kövics, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=567.97&link_type=cbs">CBS
567.97</ext-link> = PD 97/2160; from seed of <italic>Glycine max</italic>, 1997, G.
Kövics, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=100580&link_type=cbs">CBS
100580</ext-link> = PD 98/1135. <bold>The Netherlands</bold>, from <italic>Pisum
sativum</italic>, 1981, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=318.90&link_type=cbs">CBS
318.90</ext-link> = PD 81/729. <bold>U.S.A.</bold>, Minnesota, from <italic>Trifolium
pretense</italic>, 1966, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=531.66&link_type=cbs">CBS
531.66</ext-link>.</p><p><italic>Notes</italic>: <italic>Phoma sojicola</italic>, which was erected in 1999
(<xref ref-type="bibr" rid="ref129">Kövics <italic>et al.</italic>1999</xref>), has recently been synonymised with the present species,
based on morphological and genetical similarities
(<xref ref-type="bibr" rid="ref117">Irinyi <italic>et al.</italic>2009</xref>). The present study supports these findings.</p><p><italic><bold>Peyronellaea pinodes</bold></italic> (Berk. & A. Bloxam) Aveskamp,
Gruyter & Verkley, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515610&link_type=mb">MB515610</ext-link>.</p><p><italic>Basionym</italic>: <italic>Sphaeria pinodes</italic> Berk. & A. Bloxam, Ann.
Mag. Nat. Hist., Ser. III 7: 454. 1861.</p><p><list list-type="simple"><list-item><p>≡ <italic>Didymella pinodes</italic> (Berk. & A. Bloxam) Petr., Ann.
Mycol. 22(1/2): 16. 1924.</p></list-item><list-item><p>≡ <italic>Mycosphaerella pinodes</italic> (Berk. & A. Bloxam)
Vestergr<italic>.,</italic> Ann. Mycol. 10(5): 581. 1912.</p></list-item><list-item><p>= <italic>Ascochyta pinodes</italic> L.K. Jones, Bull. New York State Agric. Exp.
Sta. 547: 4. 1927.</p></list-item></list></p><p><italic>Specimens examined</italic>: <bold>Belgium</bold>, Gembloux, from <italic>Pisum
sativum</italic>, 1977, G. Sommereyns,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=525.77&link_type=cbs">CBS 525.77</ext-link>.
<bold>Iraq</bold>, Basrah province, from <italic>Pisum sativum</italic>, 1977,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=159.78&link_type=cbs">CBS 159.78</ext-link>.
<bold>Switzerland</bold>, Glarus Kanton, Filzbach, from a leaf of <italic>Primula
auricula</italic>, June 1949, E. Müller,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=285.49&link_type=cbs">CBS 285.49</ext-link>. <bold>The
Netherlands</bold>, from an unknown substrate, 1955, M.H. van Raalte,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=235.55&link_type=cbs">CBS 235.55</ext-link>.</p><p><italic><bold>Peyronellaea pomorum</bold></italic> var. <italic><bold>circinata</bold></italic>(Kusnezowa) Aveskamp, Gruyter & Verkley, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515612&link_type=mb">MB515612</ext-link>.</p><p><italic>Basionym</italic>: <italic>Peyronellaea circinata</italic> Kusnezowa, Novoste Sist.
Nizsh. Rast. 8: 189. 1971.</p><p><list list-type="simple"><list-item><p>≡ <italic>Phoma jolyana</italic> var. <italic>circinata</italic> (Kusnezowa) Boerema,
Dorenb. & Kesteren, Kew Bull. 31: 535. 1977 [1976].</p></list-item><list-item><p>≡ <italic>Phoma pomorum</italic> var. <italic>circinata</italic> (Kusnezowa)
Aveskamp, Gruyter & Verkley, Mycologia 101(3): 377. 2009.</p></list-item><list-item><p>= <italic>Peyronellaea nigricans</italic> Kusnezowa, Novoste Sist. Nizsh. Rast. 8:
191. 1971.</p></list-item></list></p><p><italic>Specimens examined</italic>: <bold>Russia</bold>, Siberia, Novosibirsk, from
<italic>Heracleum dissectum</italic>, 1963, T.T. Kusnezowa, <bold>isotype</bold> CBS H-3747,
ex-isotype culture <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=285.76&link_type=cbs">CBS
285.76</ext-link> = ATCC 26241 = IMI 176742 = VKM F-1843; Siberia,
Novosibirsk, from a leaf of <italic>Allium nutans</italic>, 1963, T.T. Kusnezowa,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=286.76&link_type=cbs">CBS 286.76</ext-link> = ATCC
26242 = IMI 176743 = VKM F-1844.</p><p><italic><bold>Peyronellaea pomorum</bold></italic> var. <italic><bold>cyanea</bold></italic> (Jooste
& Papendorf) Aveskamp, Gruyter & Verkley, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515614&link_type=mb">MB515614</ext-link>.</p><p><italic>Basionym</italic>: <italic>Phoma cyanea</italic> Jooste & Papendorf, Mycotaxon
12: 444. 1981.</p><p><list list-type="simple"><list-item><p>≡ <italic>Phoma pomorum</italic> var. <italic>cyanea</italic> (Jooste &
Papendorf) Aveskamp, Gruyter & Verkley, Mycologia 101(3): 377. 2009.</p></list-item></list></p><p><italic>Specimen examined</italic>: <bold>South Africa</bold>, Heilbron, from straw of
<italic>Triticum</italic> sp., 1972, W.J. Jooste, <bold>holotype</bold> PREM 45736,
ex-holotype culture <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=388.80&link_type=cbs">CBS
388.80</ext-link>.</p><p><italic><bold>Peyronellaea pomorum</bold></italic> var. <italic><bold>pomorum</bold></italic>(Thüm.) Aveskamp, Gruyter & Verkley, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515611&link_type=mb">MB515611</ext-link>.</p><p><italic>Basionym</italic>: <italic>Phoma pomorum</italic> var. <italic>pomorum</italic> Thüm.,
Fungi Pomicoli: 105. 1879.</p><p><italic>Specimen examined</italic>: <bold>The Netherlands</bold>, Wageningen, from
<italic>Polygonum tataricum</italic>, 1964, M.M.J. Dorenbosch, CBS H-16540, culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=539.66&link_type=cbs">CBS 539.66</ext-link> = ATCC
16791 = IMI 122266 = PD 64/914.</p><p><italic><bold>Peyronellaea protuberans</bold></italic> (Lév.) Aveskamp, Gruyter
& Verkley, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515613&link_type=mb">MB515613</ext-link>.</p><p><italic>Basionym</italic>: <italic>Phoma protuberans</italic> Lév., Ann. Sci. Nat.
Bot. III 5: 281. 1846.</p><p><italic>Specimen examined</italic>: <bold>The Netherlands</bold>, from a leaf of
<italic>Lycium halifolium</italic>, 1971,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=381.96&link_type=cbs">CBS 381.96</ext-link> = PD
71/706.</p><p><italic><bold>Peyronellaea sancta</bold></italic> <bold>(</bold>Aveskamp, Gruyter &
Verkley) Aveskamp, Gruyter & Verkley, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515615&link_type=mb">MB515615</ext-link>.</p><p><italic>Basionym</italic>: <italic>Phoma sancta</italic> Aveskamp, Gruyter & Verkley,
Mycologia 101(3): 377. 2009.</p><p><italic>Specimens examined</italic>: <bold>Argentina</bold>, from <italic>Opuntia
ficus-indica</italic>, 1997, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=644.97&link_type=cbs">CBS
644.97</ext-link>. <bold>South Africa</bold>, from dead branches of <italic>Ailanthus
altissima</italic>, Oct. 1982, C. Jansen CBS H-16332, ex-holotype culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=281.83&link_type=cbs">CBS 281.83</ext-link>. Unknown
origin, from <italic>Gleditsia triancantha</italic> culture LEV 15292.</p><p><italic><bold>Peyronellaea subglomerata</bold></italic> (Boerema, Gruyter & Noordel.)
Aveskamp, Gruyter & Verkley, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515616&link_type=mb">MB515616</ext-link>.</p><p><italic>Basionym</italic>: <italic>Phoma subglomerata</italic> Boerema, Gruyter &
Noordel., Persoonia 15(2): 204. 1993.</p><p><italic>Specimens examined</italic>: <bold>U.S.A.</bold>, North Dakota, from
<italic>Triticum</italic> sp., 1976,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=110.92&link_type=cbs">CBS 110.92</ext-link> = PD
76/1010. Unknown origin, from <italic>Zea mays</italic>, 1978, PD 78/1090.</p><p><italic><bold>Peyronellaea zeae-maydis</bold></italic> (Mukunya & Boothr.) Aveskamp,
Gruyter & Verkley, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515617&link_type=mb">MB515617</ext-link>.</p><p><italic>Basionym</italic>: <italic>Mycosphaerella zeae-maydis</italic> Mukunya &
Boothr., Phytopathology 63: 530. 1973.</p><p><list list-type="simple"><list-item><p>≡ <italic>Didymella zeae-maydis</italic> (Mukunya & Boothr.) Arx, Beih.
Nova Hedwigia 87: 288. 1987.</p></list-item></list></p><p><italic>Anamorph</italic>: <italic>Phyllosticta maydis</italic> Arny & R.R. Nelson,
Phytopathology 61: 1171. 1971.</p><p><list list-type="simple"><list-item><p>≡ <italic>Phoma zeae-maydis</italic> Punith., Mycopathologia 112(1): 50.
1990.</p></list-item></list></p><p><italic>Specimen examined</italic>: <bold>U.S.A.</bold>, Wisconsin, Hancock, from <italic>Zea
mays</italic>, June 1969, D.C. Arny, ex-holotype culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=588.69&link_type=cbs">CBS 588.69</ext-link>.</p></sec><sec><title>Group L:</title><p><italic>Phoma draconis, Ph. henningsii, Ph. plurivora</italic> and the novel
species <italic>Ph. brasiliensis</italic> cluster basally to the <italic>Epicoccum</italic>species in group M. The species clustered here, however, all lack
chlamydospores. These species do, like the chlamydospore-forming species
mentioned above, solely produce unicellular conidia, and have glabrous,
thin-walled, pseudoparenchymatous pycnidial walls composed of isodiametric
cells.</p><p><italic><bold>Phoma brasiliensis</bold></italic> Aveskamp, Gruyter & Verkley, <bold>sp.
nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515618&link_type=mb">MB515618</ext-link>.
<xref ref-type="fig" rid="fig6">Fig. 6</xref>.</p><p>Conidia cylindrica, glabra, hyalina, continua, 6–9(–10) ×
2–3(–3.5) μm, (3–)4–6(–8) guttulis parvis
praedita. Matrix conidiorum alba.</p><p><fig position="float" id="fig6"><label>Fig. 6.</label><caption><p><italic>Phoma brasiliensis</italic>(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=120105&link_type=cbs">CBS 120105</ext-link>a).
A–C. Fourteen-day-old colonies on OA (A), MEA (B) and CHA (C).
D–E. Pycnidia. F. Section of the pycnidial wall. G. Conidia. Scale bars:
D = 200 μm; E = 100 μm; F–G = 10 μm.</p></caption><graphic xlink:href="1fig6"></graphic></fig></p><p><italic>Etymology</italic>: Epithet refers to the country of origin, Brazil.</p><p><italic>Conidiomata</italic> pycnidial, mainly solitary but also confluent, globose
to irregularly shaped, glabrous, on the agar surface and immersed,
(220–)250–370(–550) ×
(150–)190–290(–320) μm. Usually with a single
inconspicuous non-papillate ostiole. <italic>Pycnidial wall</italic>pseudoparenchymatous, composed of 5–9 layers of oblong to isodiametric
cells, 18–27 μm thick. <italic>Conidiogenous cells</italic> phialidic,
hyaline, simple, smooth, globose to flask-shaped, <italic>ca</italic>. 4–5
× 3.5–4 μm. <italic>Conidia</italic> variable in size, cylindrical,
thin-walled, smooth, hyaline, aseptate 6–9(–10) ×
2–3(–3.5) μm, with (3–)4–6(–8) small polar
guttules. Conidial matrix white.</p><p><italic>Culture characteristics</italic>: Colonies on OA 50–53 mm diam after
7 d, margin regular. Aerial mycelium sparse, tufted near the centre of the
colony, white. Immersed mycelium hyaline. Abundant pycnidia produced
semi-immersed in concentric rings. Pycnidia in the outer rings pale luteous,
darkening towards the centre of the colony via buff, honey, hazel to
brown-vinaceous; reverse concolourous. Colonies on MEA 59–63 mm diam
after 7 d, margin regular. Immersed mycelium completely covered by a mycelial
mat, which is densely floccose, greenish olivaceous to greenish grey, with
elements of citrine, olivaceous black and white; reverse concolourous. Hyphae
locally containing red amorphous chrystaline material. Colonies on CHA
62–67 mm diam after 7 d, margin regular. Aerial mycelium floccose,
white. Abundant dark pycnidia are formed on the agar surface. Application of
NaOH results in a luteous discolouration of the agar, later changing to
reddish, best to be observed on OA medium.</p><p><fig position="float" id="fig7"><label>Fig. 7.</label><caption><p>Globose chlamydospores of <italic>Epicoccum</italic> spp. A–B. <italic>E.
nigrum</italic> (<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=173.73&link_type=cbs">CBS
173.73</ext-link>). C–E. <italic>E. sorghi</italic>(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=246.60&link_type=cbs">CBS 246.60</ext-link>).
F–H. <italic>E. pimprinum</italic>(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=179.80&link_type=cbs">CBS 179.80</ext-link>). Scale
bars: A–B = 50 μm; C–H = 20 μm.</p></caption><graphic xlink:href="1fig7"></graphic></fig></p><p><italic>Specimen examined</italic>: <bold>Brazil</bold>, from <italic>Amaranthus</italic> sp.,
Nov. 2007, E. Rosskopf, <bold>holotype designated here</bold> CBS H-20235,
ex-holotype culture <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=120105&link_type=cbs">CBS
120105</ext-link>.</p><p><italic>Notes</italic>: This species is thus far only known from a single isolate
from a wild <italic>Amaranthus</italic> sp. in Brazil. According to Boerema <italic>et
al.</italic> (<xref ref-type="bibr" rid="ref55">2004</xref>), no other
<italic>Phoma</italic> species have been recorded from the same host.</p><p><italic><bold>Phoma draconis</bold></italic> (Berk. ex Cooke) Boerema, Verslagen Meded.
Plziektenk. Dienst Wageningen 159 (Jaarboek 1982): 24. 1983.</p><p><italic>Basionym</italic>: <italic>Phyllosticta draconis</italic> Berk. ex Cooke, Grevillea
19: 8. 1891.</p><p><italic>Specimen examined</italic>: <bold>Rwanda</bold>, from a leaf of <italic>Dracaena</italic>sp., Jan. 1982, G.H. Boerema, CBS H-16207, culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=186.83&link_type=cbs">CBS 186.83</ext-link> = PD
82/47.</p><p><italic><bold>Phoma henningsii</bold></italic> Sacc., Syll. Fung. 10: 139. 1892.</p><p><italic>Specimen examined</italic>: <bold>Kenya</bold>, Maguga, from the bark of
<italic>Acacia mearnsii</italic>, June 1992, T.W. Olembo, CBS H-16354, culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=104.80&link_type=cbs">CBS 104.80</ext-link> = PD
74/1017.</p><p><italic><bold>Phoma plurivora</bold></italic> P.R. Johnst., New Zealand J. Bot. 19(2):
181. 1981.</p><p><italic>Specimens examined</italic>: <bold>Australia</bold>, from <italic>Medicago
sativa</italic>, 1975, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=248.93&link_type=cbs">CBS
248.93</ext-link> = PD 95/907. <bold>New Zealand</bold>, Auckland, Mt Albert, from
a leaf of <italic>Setaria sp</italic>., Feb. 1979, P.R. Johnston, CBS H-7624,
ex-isotype culture <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=558.81&link_type=cbs">CBS
558.81</ext-link> = PDDCC 6873.</p></sec><sec><title>Group M – Epicoccum:</title><p>This group (BPP = 1.00, RBS = 66 %) comprises three species that are
accommodated in the section <italic>Peyronellaea</italic>. The <italic>Peyronellaea</italic>species in this group, <italic>Ph. sorghina, Ph. pimprina</italic> and <italic>Epicoccum
nigrum</italic> (chlamydospore-based synanamorph of <italic>Ph. epicoccina</italic>;
Arenal <italic>et al.</italic> <xref ref-type="bibr" rid="ref12">2000</xref>,
<xref ref-type="bibr" rid="ref13">2004</xref>) are characterised by the
production of botryoid or epicoccoid chlamydospores, in contrast to the
species in group K, which produce alternarioid dictyochlamydospores. The
distinct morphology and phylogenetic position justify the recombination into a
separate genus. As the oldest generic name in this clade is
<italic>Epicoccum</italic>, new combinations for <italic>Ph. pimprina</italic> and <italic>Ph.
sorghina</italic> are proposed below.</p><p><italic><bold>Epicoccum</bold></italic> Link, Mag. Gesell. Naturf. Freunde Berlin 7: 32.
1815, <bold>emend.</bold> Aveskamp, Gruyter & Verkley.
<xref ref-type="fig" rid="fig7">Fig. 7</xref>.</p><p><italic>Conidiomata</italic> pycnidial, globose to subglobose, measuring
50–250 μm diam, on agar surface or immersed, mostly solitary but
incidentally confluent. <italic>Ostioles</italic> papillate or on pronounced necks.
<italic>Pycnidial wall</italic> pseudoparenchymatous, counting 2–8 cell layers
of which the outer 1–3 are brown-olivaceous pigmented. <italic>Conidiogenous
cells</italic> phialidic, hyaline, simple, smooth, ampulliform, <italic>ca</italic>.
3–7 × 3–7 μm. <italic>Conidia</italic> variable in shape,
initially hyaline but in later stages a slight brownish pigmentation may be
found, thin-walled, smooth, always aseptate 3–8.5(–10) ×
1.5–4(–4.5) μm. <italic>Chlamydospores</italic> unicellular or
multicellular, intercalary or terminal, smooth, verrucose or incidentally
tuberculate, subhyaline to dark brown, where unicellular globose, measuring
5–15 μm diam, where multicellular globose or irregular shaped,
smooth, verrucose or incidentally tuberculate, measuring 8–35 μm.</p><p><italic>Type species</italic>: <italic>Epicoccum nigrum</italic> Link.</p><p><italic><bold>Epicoccum nigrum</bold></italic> Link, Mag. Gesell. Naturf. Freunde Berlin
7: 32. 1815.</p><p><list list-type="simple"><list-item><p>≡ <italic>Phoma epicoccina</italic> Punith., M.C. Tulloch & C.M. Leach,
Trans. Brit. Mycol. Soc. 59(2): 341 (1972).</p></list-item></list></p><p><italic>Specimens examined</italic>: <bold>Germany</bold>, Berlin, from soil, 1985, H.J.
Halfmann, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=505.85&link_type=cbs">CBS
505.85</ext-link>. <bold>The Netherlands</bold>, Geleen, from human toe nail, Dec.
1981, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=125.82&link_type=cbs">CBS 125.82</ext-link> =
IMI 331914 = CECT 20044; Randwijk, from <italic>Malus</italic> sp., J. Köhl,
2003, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=115825&link_type=cbs">CBS 115825</ext-link>.
<bold>U.S.A.</bold>, Oregon, from seeds of <italic>Dactylis glomerata</italic>, 1967,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=173.73&link_type=cbs">CBS 173.73</ext-link> = ATCC
24428 = IMI 164070.</p><p><italic><bold>Epicoccum pimprinum</bold></italic> (P.N. Mathur, S.K. Menon & Thirum.)
Aveskamp, Gruyter & Verkley, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515619&link_type=mb">MB515619</ext-link>.</p><p><italic>Basionym</italic>: <italic>Phoma pimprina</italic> P.N. Mathur, S.K. Menon &
Thirum., Sydowia 13: 146. 1959.</p><p><italic>Specimens examined</italic>: <bold>India</bold>, Poona, Pimpri, from soil, Mar.
1959, S.K. Menon, ex-isotype culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=246.60&link_type=cbs">CBS 246.60</ext-link> = ATCC
22237 = ATCC 16652 = IMI 81601; from soil, 1977, PD 77/1028.</p><p><italic><bold>Epicoccum sorghi</bold></italic> (Sacc.) Aveskamp, Gruyter & Verkley,
<bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515620&link_type=mb">MB515620</ext-link>.</p><p><italic>Basionym</italic>: <italic>Phyllosticta sorghina</italic> Sacc., <italic>Michelia</italic><bold>1</bold> (2): 140. 1878.</p><p><list list-type="simple"><list-item><p>≡ <italic>Phoma sorghina</italic> (Sacc.) Boerema, Dorenb. & Kesteren,
Persoonia 7(2): 139. 1972.</p></list-item></list></p><p>For a complete synonymy see Boerema <italic>et al.</italic>(<xref ref-type="bibr" rid="ref48">1977</xref>).</p><p><italic>Specimens examined</italic>: <bold>France</bold>, Antibes, from a twig of
<italic>Citrus</italic> sp., 1966, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=627.68&link_type=cbs">CBS
627.68</ext-link> = PD 66/926. <bold>Guinea-Bissau</bold>, Gacheu Région,
from <italic>Oryza sativa</italic>, Oct. 1978,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=181.81&link_type=cbs">CBS 181.81</ext-link>.
<bold>India</bold>, from a fruit of <italic>Coffea</italic> sp., July 1968, C.V.
Subramanian, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=846.68&link_type=cbs">CBS
846.68</ext-link>; Jabalpur, from <italic>Panicum miliare</italic>, Jan. 1972, D.
Sharma, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=293.72&link_type=cbs">CBS 293.72</ext-link>.
<bold>Martinique</bold>, from a leaf of <italic>Lycopersicon esculentum</italic>, June 1989,
B. Hostachy, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=301.89&link_type=cbs">CBS
301.89</ext-link>. <bold>Papua New Guinea</bold>, from <italic>Stellaria</italic> sp., A.
Aptroot, Oct. 1995, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=886.95&link_type=cbs">CBS
886.95</ext-link>; Central Province, Varirata National Park near Port
Moresby, from soil, A. Aptroot, Oct. 1995,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=986.95&link_type=cbs">CBS 986.95</ext-link>.
<bold>Puerto Rico</bold>, Mayaguez, from <italic>Sorghum vulgare</italic>, Apr. 1976, R.
Alconera, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=179.80&link_type=cbs">CBS
179.80</ext-link>= PD 76/1018. <bold>South Africa</bold>, Potchefstroom, from a
leaf of <italic>Zea mays</italic>, Nov. 1978, W.J. Jooste,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=180.80&link_type=cbs">CBS 180.80</ext-link> = PD
78/1100.</p><p><italic>Notes</italic>: The strains that were previously accommodated in <italic>Ph.
sorghina</italic> are morphologically and phylogenetically highly diverse
(<xref ref-type="bibr" rid="ref19">Aveskamp <italic>et al.</italic>2009a</xref>,
<xref ref-type="bibr" rid="ref161">Pažoutová
2009</xref>), and probably represent multiple species. These species
were, however, not treated in the present study.</p></sec><sec><title>Group N – Boeremia gen. nov.:</title><p>This group represents species that are morphologically similar to what is
currently known as <italic>Ph. exigua</italic>. Group N is a well-defined clade (BPP =
1.00, RBS = 100 %) and comprises all taxa that were previously recognised as
separate <italic>Ph. exigua</italic> varieties by Abeln <italic>et al.</italic>(<xref ref-type="bibr" rid="ref8">2002</xref>). <italic>Phoma foveata</italic>and <italic>Ph. sambuci-nigrae</italic> are embedded here as well, two species that
previously were known as varieties of <italic>Ph. exigua</italic>, but were elevated
to species rank due to their phytopathological relevance (<italic>Ph.
foveata</italic>, <xref ref-type="bibr" rid="ref51">Boerema <italic>et al.</italic>1987</xref>) or distinct physiological characters (<italic>Ph.
sambuci-nigrae</italic>, <xref ref-type="bibr" rid="ref137">Monte <italic>et
al.</italic> 1991</xref>). As already noted by Aveskamp <italic>et al.</italic>(<xref ref-type="bibr" rid="ref20">2009b</xref>) also <italic>Ph. telephii,
Ph. strasseri</italic> and <italic>Ph. lycopersici</italic> are closely related. This
study also reveals the close relationship with <italic>Ph. tarda</italic>, a pathogen
of coffee. <italic>Phoma hedericola</italic>, a frequently occurring causal agent of
leaf spots on poison ivy (<italic>Hedera helix</italic>) and <italic>Ph. crinicola</italic>, a
pathogen of <italic>Amaryllidaceae</italic> are embedded in this clade. In contrast to
the other species in this clade, which are linked to <italic>Phoma</italic> section
<italic>Phyllostictoides, Ph. hedericola</italic> and <italic>Ph. crinicola</italic> are
associated with <italic>Phoma</italic> section <italic>Phoma</italic>, due to the absence of
septate conidia (<xref ref-type="bibr" rid="ref96">De Gruyter &
Noordeloos 1992</xref>, <xref ref-type="bibr" rid="ref97">De Gruyter
<italic>et al.</italic> 1993</xref>). The sequence data of
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=172.34&link_type=cbs">CBS 172.34</ext-link>, a
strain recorded as <italic>Dothiorella ulmi,</italic> appeared to be genetically
identical to <italic>Ph. exigua,</italic> as was already noted by De Gruyter <italic>et
al.</italic> (<xref ref-type="bibr" rid="ref100">2009</xref>). Based on
morphological studies of other strains, <italic>Dothiorella ulmi</italic> was
suggested to be recombined into <italic>Plectophomella</italic>(<xref ref-type="bibr" rid="ref179">Redfern & Sutton 1981</xref>),
a genus that is linked to the Pezizomycotina. Morphological features of the
present strain appeared to be similar to <italic>Ph. exigua</italic>, suggesting that
this strain was probably preserved under an incorrect name, and actually
belongs to <italic>Ph. exigua</italic> var. <italic>populi</italic>.</p><p>Of the species within this clade, a teleomorph is only named in <italic>Ph.
lycopersici</italic> (<italic>Didymella lycopersici</italic>), although Stewart
(<xref ref-type="bibr" rid="ref212">1957</xref>) has reported the
existence of pseudothecia of <italic>Ph. tarda</italic> in nature, a finding that also
has been reported by Salgado <italic>et al.</italic>(<xref ref-type="bibr" rid="ref190">2007</xref>). This contradicts with
the fact that none of the varieties embedded in the <italic>Ph. exigua</italic> has
been found in association with a teleomorph thus far.</p><p>For further delineation of this clade, a comparison of actin gene sequences
is proposed (<xref ref-type="bibr" rid="ref20">Aveskamp <italic>et al</italic>.
2009b</xref>), although not all species and varieties in this complex can
be recognised using this gene only. Thus far the varieties of <italic>Ph.
exigua</italic> could only be delineated using two fingerprint techniques:
Amplified Fragment Length Polymorphism (AFLP,
<xref ref-type="bibr" rid="ref8">Abeln <italic>et al.</italic> 2002</xref>) and
DAF (DNA Amplification Fingerprinting) using mini-hairpin primers
(<xref ref-type="bibr" rid="ref20">Aveskamp <italic>et al.</italic>2009b</xref>). Based on this latter technique Aveskamp <italic>et al.</italic>(<xref ref-type="bibr" rid="ref20">2009b</xref>) recognised two
varieties within <italic>Ph. exigua</italic> that had not been described before. These
two infraspecific taxa, var. <italic>gilvescens</italic> and var.
<italic>pseudolilacis</italic> are treated and described below.</p><p>Based on the phylogenetic reconstruction obtained here, the taxa previously
known as <italic>Ph. exigua</italic> var. <italic>noackiana</italic> and <italic>Ph. exigua</italic>var. <italic>diversispora</italic> cluster in a distinct clade from the other
varieties in this complex, and are elevated to species level here. Also actin
sequence data and DAF analysis (<xref ref-type="bibr" rid="ref20">Aveskamp
<italic>et al</italic>. 2009b</xref>), AFLP data
(<xref ref-type="bibr" rid="ref8">Abeln <italic>et al.</italic> 2002</xref>)
reveal a basal topology of these species compared to <italic>Ph. exigua</italic>.
Morphological data obtained by Van der Aa (2000) also suggest that these
species are not completely fitting in the <italic>Ph. exigua</italic> concept.</p><p>The species and varieties in this clade differ from other <italic>Phoma</italic>taxa based on their ostiole morphology. In contrast to other species, which
have a smoothly lined ostiole, the taxa present in this clade have distinct
hyaline cells lining their ostiolar openings
(<xref ref-type="fig" rid="fig8">Fig. 8A</xref>). In addition, these
species, with the exception of <italic>Ph. hedericola</italic>, produce septate
conidia in addition to the regular aseptate ones, although in general the
septate conidia are produced in smaller numbers in culture than on the host.
These conidia are mostly 1-septate, as only in <italic>Ph. exigua</italic>incidentally multiseptate conidia occur, and are often only slightly larger
than the regular aseptate ones (<xref ref-type="fig" rid="fig8">Fig.
8C</xref>). Due to the morphological and genetic distinctiveness, we
propose a new generic name for the taxa in this clade.</p><p><italic><bold>Boeremia</bold></italic> Aveskamp, Gruyter & Verkley, <bold>gen. nov.</bold>MycoBank <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515621&link_type=mb">MB515621</ext-link>.
<xref ref-type="fig" rid="fig8">Fig. 8</xref>.</p><p>Conidiomata pycnidialia, plerumque globosa vel subglobosa, glabra vel
eminentiis sparsis hypharum vestita, superficialia vel in agaro immersa,
solitaria vel confluentia, 75–370 μm diam. Ostiola papillata vel
epapillata, tempore maturitatis interne cellulis hyalinis papillatis. Paries
pycnidii pseudoparenchymatus, e 2–8 stratis cellularum compositus,
extima 1–3 strata brunnea. Cellulae conidiogenae phialidicae, hyalinae,
glabrae, ampulliformes vel dolliiformes, <italic>ca.</italic> 3–7.5 ×
3–6.5 μm. Conidia hyalina, tenuitunicata, glabra, plerumque continua,
2.5–12 × 2–4 μm, et interdum uni- vel biseptata, usque 15
× 5 μm.</p><p><italic>Conidiomata</italic> pycnidial conidiomata variable in shape and size,
mostly globose to subglobose, glabrous or with few mycelial outgrowths, on
agar surface or immersed, solitary or confluent, measuring 75–370 μm
diam. <italic>Ostioles</italic> 1–2(–3), non-pappillate or pappillate,
lined internally with a pappillate hyaline cells when mature. <italic>Pycnidial
wall</italic> pseudoparenchymatous, counting 2–8 cell layers of which the
outer 1–3 are brown pigmented. <italic>Conidiogenous cells</italic> phialidic,
hyaline, simple, smooth, ampulliform to doliiform, <italic>ca</italic>. 3–7.5
× 3–6.5 μm. <italic>Conidia</italic> variable in shape, hyaline,
thin-walled, smooth, mainly aseptate, 2.5–12 × 2–4 μm,
but regularly 1(–2)-septate conidia may be found which measure up to 15
× 5 μm. <italic>Pseudothecia</italic>, only rarely recorded in one species
<italic>in vivo</italic>, subglobose, up to 300 μm diam. <italic>Asci</italic> cylindrical
or subclavate, measuring 50–95 × 6–10 μm, always
8-spored, biseriate. <italic>Ascospores</italic> ellipsoid, measuring 12–18
× 5–6 μm, uniseptate.</p><p><italic>Type species</italic>: <italic>Boeremia exigua</italic> (Desm.) Aveskamp, Gruyter
& Verkley</p><p><italic>Etymology</italic>: Named after Gerhard H. Boerema, who made great
contributions to our understanding of the taxonomy of phomoid fungi.</p><p><fig position="float" id="fig8"><label>Fig. 8.</label><caption><p><italic>Boeremia</italic> gen. nov. A. Ostiole configuration of <italic>B. exigua</italic>var. <italic>exigua</italic> (<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=431.74&link_type=cbs">CBS
431.74</ext-link>). B. Pycnidial wall and conidiogenous cells of <italic>B.
telephii</italic> (<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=760.73&link_type=cbs">CBS
760.73</ext-link>) C. Aseptate and septate conidia of <italic>B. lycopersici</italic>(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=378.67&link_type=cbs">CBS 378.67</ext-link>). Scale
bars: A = 20 μm; B–C = 10 μm.</p></caption><graphic xlink:href="1fig8"></graphic></fig></p><p><italic><bold>Boeremia crinicola</bold></italic> (Siemasko) Aveskamp, Gruyter &
Verkley, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515622&link_type=mb">MB515622</ext-link>.</p><p><italic>Basionym</italic>: <italic>Phyllosticta crinicola</italic> Siemasko, Acta Soc. Bot.
Poloniae 1: 22. 1923.</p><p><list list-type="simple"><list-item><p>≡ <italic>Phoma crinicola</italic> (Siemasko) Boerema apud Boerema &
Dorenbosch, Verslagen Meded. Plziektenk. Dienst Wageningen 153: 18.1979.</p></list-item></list></p><p><italic>Specimens examined</italic>: <bold>The Netherlands</bold>, Haarlem, from a bulb
of <italic>Crinum powellii</italic>, Mar. 1976, G.H. Boerema, CBS H-16198,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=109.79&link_type=cbs">CBS 109.79</ext-link> = PD
77/747; Alkmaar, from a bulb of <italic>Crinum</italic> sp., 1970, G.H. Boerema,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=118.93&link_type=cbs">CBS 118.93</ext-link> = PD
70/195.</p><p><italic><bold>Boeremia diversispora</bold></italic> (Bubák) Aveskamp, Gruyter
& Verkley, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515623&link_type=mb">MB515623</ext-link>.</p><p><italic>Basionym</italic>: <italic>Phoma diversispora</italic> Bubák, Oest. Bot. Z.
55: 78. 1905</p><p><list list-type="simple"><list-item><p>≡ <italic>Phoma exigua</italic> var. <italic>diversispora</italic> (Bubák)
Boerema apud Boerema & van Kesteren, Gewasbescherming 11: 122. 1980</p></list-item></list></p><p>For a complete description see Boerema <italic>et al.</italic>(<xref ref-type="bibr" rid="ref49">1981a</xref>,
<xref ref-type="bibr" rid="ref55">2004</xref>), and Van der Aa <italic>et
al.</italic> (<xref ref-type="bibr" rid="ref7">2000</xref>).</p><p><italic>Specimens examined</italic>: <bold>Brazil</bold>, leaf of <italic>Phaseolus</italic>, F.
Noack, <bold>holotype</bold> B. <bold>Kenya</bold>, from a pod of <italic>Phaseolus
vulgaris</italic>, 1979, G.H. Boerema, <bold>epitype designated here</bold> CBS H-16308,
ex-epitype culture <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=102.80&link_type=cbs">CBS
102.80</ext-link> = CECT 20049 = IMI 331907 = PD 79/61. <bold>The
Netherlands</bold>, near Tilburg, from <italic>Phaseolus vulgaris</italic>, 1979, J. de
Gruyter, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=101194&link_type=cbs">CBS 101194</ext-link>= PD 79/687 = IMI 373349.</p><p><italic>Notes</italic>: <italic>Phoma diversispora</italic> was originally described by
Bubák as a pathogen of cowpea (<italic>Vigna unguiculata</italic>) causing
Black Node Disease (<xref ref-type="bibr" rid="ref7">Van der Aa <italic>et
al.</italic> 2000</xref>), but was later classified as a variety of <italic>Ph.
exigua</italic> by Boerema & Van Kesteren
(<xref ref-type="bibr" rid="ref42">1980</xref>) and Boerema <italic>et
al</italic>. (<xref ref-type="bibr" rid="ref49">1981a</xref>), on basis of
its morphology. The present study, however, revealed the <italic>B. exigua</italic>varieties to be phylogenetically distinct from the present species, which
justifies re-establishment of the taxon as separate species in the genus
<italic>Boeremia</italic>. The present species is closely related to <italic>B.
noackiana</italic>, formerly known as <italic>Ph. exigua</italic> var. <italic>noackiana</italic>(see below).</p><p><italic><bold>Boeremia exigua</bold></italic> var. <italic><bold>coffeae</bold></italic> (Henn.)
Aveskamp, Gruyter & Verkley, <bold>stat. et comb. nov.</bold>
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515632&link_type=mb">MB515632</ext-link>.</p><p><italic>Basionym</italic>: <italic>Ascochyta coffeae</italic> Henn., Hedwigia 41: 307.
1902; not <italic>Phoma coffeae</italic> Delacr., Bull. Soc. Mycol. France 13: 122.
1897.</p><p><list list-type="simple"><list-item><p>= <italic>Ascochyta tarda</italic> R.B. Stewart, Mycologia 49: 430. 1957.</p></list-item><list-item><p>≡ <italic>Phoma tarda</italic> (R.B. Stewart) H. Verm., Coffee Berry Dis.
Kenya: 14. 1979.</p></list-item></list></p><p>For a complete description see De Gruyter <italic>et al.</italic>(<xref ref-type="bibr" rid="ref99">2002</xref>).</p><p><italic>Specimens examined</italic>: <bold>Brazil</bold>, Patrocínio, from leaf of
<italic>Coffea arabica</italic>, L.H. Pfenning,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=119730&link_type=cbs">CBS 119730</ext-link>.
<bold>Cameroon</bold>, Bemenda, from <italic>Coffea arabica</italic>,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=109183&link_type=cbs">CBS 109183</ext-link> = PD
2000/10506 = IMI 300060.</p><p><italic>Notes</italic>: <italic>Boeremia exigua</italic> var. <italic>coffeae</italic> was
originally described from leaves of coffee plants (<italic>Coffea arabica</italic>,
<xref ref-type="bibr" rid="ref212">Stewart 1957</xref>) as
<italic>Ascochyta coffeae</italic> and <italic>A. tarda</italic>. The observed late
euseptation in this species proved to be a character common for <italic>Phoma</italic>species accommodated in section <italic>Phyllostictoides</italic>, leading to a
recombination into <italic>Phoma</italic>, as <italic>Ph. tarda</italic>. Phylogenetic results
obtained in the present study reveal genetic similarity between the present
species and the <italic>B. exigua</italic> species complex. The cultures of <italic>B.
exigua</italic> varieties are somewhat slower growing than those of the present
species, which completely covers the agar surface (90 mm diam) within 7 d. The
pycnidia of <italic>B. exigua</italic> var. <italic>tarda</italic> may grown to up to 255
μm (<xref ref-type="bibr" rid="ref99">De Gruyter <italic>et al.</italic>2002</xref>), but other micromorphological characters fit within the
scope of <italic>B. exigua</italic> as described for Ph. exigua by Van der Aa <italic>et
al.</italic> (<xref ref-type="bibr" rid="ref7">2000</xref>) and De Gruyter
<italic>et al.</italic> (<xref ref-type="bibr" rid="ref99">2002</xref>). It is
concluded, therefore, that <italic>Ph. tarda</italic> should be reduced to a variety
of the <italic>B. exigua</italic>. Multiple <italic>Phoma</italic> species have been found in
association with <italic>Coffea arabica</italic>, such as <italic>Ph. coffeae-arabicae,
Ph. coffeicola, Ph. coffeiphila, Ph. costarricensis, Ph. excelsa</italic>, and
<italic>Ph. pereupyrena</italic> (<xref ref-type="bibr" rid="ref189">Saccas
1981</xref>, <xref ref-type="bibr" rid="ref19">Aveskamp <italic>et al</italic>.
2009a</xref>). None of these species however matches the description that
is applied to taxa in the <italic>B. exigua</italic> complex.</p><p><italic><bold>Boeremia exigua</bold></italic> var. <italic><bold>exigua</bold></italic> (Desm.)
Aveskamp, Gruyter & Verkley, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515624&link_type=mb">MB515624</ext-link>.</p><p><italic>Basionym</italic>: <italic>Phoma exigua</italic> Desm., Ann. Sci. Nat. Bot. III 11:
282. 1849.</p><p><italic>Specimens examined</italic>: <bold>Germany</bold>, Artern, from <italic>Foeniculum
vulgare</italic>, Apr. 1984, S. Petzoldt,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=391.84&link_type=cbs">CBS 391.84</ext-link>. <bold>The
Netherlands</bold>, from a tuber of <italic>Solanum tuberosum</italic>, 1928,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=236.28&link_type=cbs">CBS 236.28</ext-link>;
Emmeloord, from a tuber of <italic>Solanum tuberosum</italic>, 1974, G.H. Boerema,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=431.74&link_type=cbs">CBS 431.74</ext-link> = PD
74/2447; Emmeloord, from <italic>Cichorium intybus</italic>, 1979, G.H. Boerema,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=101150&link_type=cbs">CBS 101150</ext-link> = PD
79/118; Ommen, from <italic>Digitalis</italic> sp., 1990, J. de Gruyter,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=101152&link_type=cbs">CBS 101152</ext-link> = PD
90/835-3.</p><p><italic><bold>Boeremia exigua</bold></italic> var. <italic><bold>forsythiae</bold></italic> (Sacc.)
Aveskamp, Gruyter & Verkley, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515625&link_type=mb">MB515625</ext-link>.</p><p><italic>Basionym</italic>: <italic>Phyllosticta forsythiae</italic> Sacc., Michelia 1(1):
93. 1997.</p><p><list list-type="simple"><list-item><p>≡ <italic>Phoma exigua</italic> var. <italic>forsythiae</italic> (Sacc.) Aa, Boerema
& Gruyter, Persoonia 17: 452. 2000.</p></list-item></list></p><p><italic>Specimens examined</italic>: <bold>The Netherlands</bold>, from
<italic>Forsythia</italic> sp., 1992, J. de Gruyter,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=101213&link_type=cbs">CBS 101213</ext-link> = PD
92/959; from <italic>Forsythia</italic> sp., 1995, J. de Gruyter,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=101197&link_type=cbs">CBS 101197</ext-link>= PD
95/721.</p><p><italic><bold>Boeremia exigua</bold></italic> var. <italic><bold>gilvescens</bold></italic> Aveskamp,
Gruyter & Verkley, <bold>var. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515626&link_type=mb">MB515626</ext-link>.
<xref ref-type="fig" rid="fig9">Fig. 9</xref>.</p><p>Varietas Phomae exiguae similis, sed matrix conidiorum flavida vel luteola.
In agaro et in mycelio aereo catenis cellularum inflatarum
(11.5–)12.5–27.5(–31) ×
(5.5–)7.5–14.5(–18) μm.</p><p><italic>Etymology</italic>: Varietal name refers to the yellow conidial matrix,
which distinguishes this variety.</p><p><fig position="float" id="fig9"><label>Fig. 9.</label><caption><p><italic>Boeremia exigua</italic> var. <italic>gilvescens</italic>(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=101150&link_type=cbs">CBS 101150</ext-link>).
A–C. Fourteen-day-old colonies on OA (A), MEA (B) and CHA (C). D.
Pycnidia. E. Chains of wollen cells. F. Conidia. Scale bars: D = 100 μm; E
= 100 μm; F = 10 μm.</p></caption><graphic xlink:href="1fig9"></graphic></fig></p><p><italic>Culture characteristics</italic>: Colonies on OA 75–80 mm diam after
7 d, margin regular or irregular. Immersed mycelium sparsely visible due to
coverage by the aerial mycelium, hyaline or black to greenish olivaceous, with
many pycnidia; reverse mouse-grey to olivaceous. Colonies on MEA 70–75
mm diam after 7 d, margin regular or irregular. Immersed mycelium completely
covered by a compact aerial mat, which is smoke-grey with some mouse-grey
zones; reverse black. Colonies on CHA at least (75–)80 mm diam after 7
d, but often the agar surface is completely covered, margin regular or
somewhat crenate. Immersed mycelium completely covered by a compact smoke-grey
mat of aerial mycelium, or, in some zones floccose, olivaceous with white
tufts; reverse shows a dendritic leaden-black zone around the colony centre,
with black zones near the colony border. Application of NaOH did not have any
effect.</p><p>Pycnidial and conidial shapes and sizes fit within the <italic>Ph. exigua</italic>species concept. Conidial matrix yellowish or pale luteous. Brown pigmented
swollen cells occur in chains in the agar and in the aerial mycelium,
measuring (11.5–)12.5–27.5(–31) ×
(5.5–)7.5–14.5(–18) μm.</p><p><italic>Specimens examined</italic>: <bold>Philippines</bold>, from <italic>Solanum
tuberosum</italic>,1990, L.J. Turkensteen,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=101156&link_type=cbs">CBS 101156</ext-link> = PD
90/731; <bold>The Netherlands</bold>; from a graft of <italic>Ulmus</italic>, 1961, H.M.
Heybroek, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=373.61&link_type=cbs">CBS
373.61</ext-link>; Baarn, from leaves of <italic>Dactylis purpurea</italic>, 1970,
H.A. van der Aa, <bold>holotype designated here</bold> CBS H-16281, culture
ex-holotype <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=761.70&link_type=cbs">CBS
761.70</ext-link>; Lisse, from <italic>Dahlia</italic>, 1982, J. de Gruyter,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=101151&link_type=cbs">CBS 101151</ext-link> = PD
82/1022.</p><p><italic>Notes</italic>: This novel variety of <italic>B. exigua</italic>, distinguished
from other <italic>B. exigua</italic> varieties on basis of DAF analysis
(<xref ref-type="bibr" rid="ref20">Aveskamp <italic>et al.</italic>2009b</xref>), is closely related to <italic>B. exigua</italic> var.
<italic>exigua</italic>, but different in the colour of its conidial matrix
(yellowish) and absence of a positive reaction to NaOH. This variety may be
identical to <italic>Ph. exigua</italic> var. <italic>inoxydabilis</italic> Boerema &
Vegh, but as the type culture has been lost
(<xref ref-type="bibr" rid="ref7">Van der Aa <italic>et al.</italic>2000</xref>) a proper comparison of the varieties cannot be made.
Additionally, <italic>Ph. exigua</italic> var. <italic>inoxydabilis</italic> was originally
only known from periwinkle (<italic>Vinca minor</italic>,
<xref ref-type="bibr" rid="ref227">Vegh <italic>et al.</italic> 1974</xref>),
whereas the strains associated to the present taxon are isolated from a wide
range of host plants.</p><p><italic><bold>Boeremia exigua</bold></italic> var. <italic><bold>heteromorpha</bold></italic> (Schulzer
& Sacc.) Aveskamp, Gruyter & Verkley, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515627&link_type=mb">MB515627</ext-link>.</p><p><italic>Basionym</italic>: <italic>Phoma heteromorpha</italic> Schulzer & Sacc.,
Hedwigia 23: 107. 1884.</p><p><list list-type="simple"><list-item><p>≡ <italic>Phoma exigua</italic> var. <italic>heteromorpha</italic> (Schulzer &
Sacc.) Noordel. & Boerema, Verslagen Meded. Plziektenk. Dienst Wageningen
166: 109.1989.</p></list-item></list></p><p><italic>Specimens examined</italic>: <bold>France</bold>, Antibes, from <italic>Nerium
oleander</italic>, 1979, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=101196&link_type=cbs">CBS
101196</ext-link> = PD 79/176. <bold>Italy</bold>, Perugia, from <italic>Nerium
oleander</italic>, 1994, A. Zazzerini,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=443.94&link_type=cbs">CBS 443.94</ext-link>.</p><p><italic><bold>Boeremia exigua</bold></italic> var. <italic><bold>lilacis</bold></italic> (Sacc.)
Aveskamp, Gruyter & Verkley, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515628&link_type=mb">MB515628</ext-link>.</p><p><italic>Basionym</italic>: <italic>Phoma herbarum</italic> f. <italic>lilacis</italic> Sacc.,
Michelia 2(1): 93. 1880.</p><p><italic>Specimen examined</italic>: <bold>The Netherlands</bold>, Wageningen, from a twig
of <italic>Syringa vulgaris</italic>, June 1976, G.H. Boerema, CBS H-163131, culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=569.79&link_type=cbs">CBS 569.79</ext-link> = PD
72/741.</p><p><italic>Notes</italic>: Although in the present study this variety clusters outside
the <italic>B. exigua</italic> cluster, it is phylogenetic affiliation is ambiguous.
In previous studies in which fingerprint markers and actin sequences were
applied to delineate this species complex
(<xref ref-type="bibr" rid="ref8">Abeln <italic>et al.</italic> 2002</xref>,
<xref ref-type="bibr" rid="ref20">Aveskamp <italic>et al.</italic>2009b</xref>) the present taxon clusters within <italic>Ph. exigua</italic>, and
is therefore recombined as <italic>B. exigua</italic> var. <italic>lilacis</italic>. Further
analysis of this complex is, however, advocated.</p><p><fig position="float" id="fig10"><label>Fig. 10.</label><caption><p><italic>Boeremia exigua</italic> var. <italic>pseudolilacis</italic>(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=101207&link_type=cbs">CBS 101207</ext-link>).
A–C. Fourteen-day-old colonies on OA (A), MEA (B) and CHA (C). D.
Pycnidia. E. section of young pycnidium. F. Conidia. Scale bars: D = 100
μm; E = 20 μm; F = 5 μm.</p></caption><graphic xlink:href="1fig10"></graphic></fig></p><p><italic><bold>Boeremia exigua</bold></italic> var. <italic><bold>linicola</bold></italic> (Naumov &
Vassiljevsky) Aveskamp, Gruyter & Verkley, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515629&link_type=mb">MB515629</ext-link>.</p><p><italic>Basionym</italic>: <italic>Ascochyta linicola</italic> Naumov & Vassiljevsky,
Mater. Micol. Fitopatol. 5: 3. 1926.</p><p><italic>Specimens examined</italic>: <bold>The Netherlands</bold>, Zierikzee, from
<italic>Linum usitatissimum</italic>, 1928, H.A. Diddens,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=114.28&link_type=cbs">CBS 114.28</ext-link>;
Flevoland, from a stem of <italic>Linum usitatissimum</italic>, 1976, G.H. Boerema,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=116.76&link_type=cbs">CBS 116.76</ext-link> = ATCC
32332 = CECT 20022 = CECT 20023 = IMI 197074 = PD 75/544.</p><p><italic><bold>Boeremia exigua</bold></italic> var. <italic><bold>populi</bold></italic> (Gruyter &
P. Scheer) Aveskamp, Gruyter & Verkley, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515630&link_type=mb">MB515630</ext-link></p><p><italic>Basionym</italic>: <italic>Phoma exigua</italic> var. <italic>populi</italic> Gruyter &
P. Scheer, J. Phytopathol. 146(8–9): 413. 1998.</p><p><italic>Specimens examined</italic>: <bold>The Netherlands</bold>, Deil, from a twig of
<italic>Populus</italic> X <italic>euramericana</italic> cv. Robusta, Feb. 1993, A.J.P. Oort,
<bold>holotype</bold> L 995.263.325, ex-holotype culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=100167&link_type=cbs">CBS 100167</ext-link> = PD
93/217; Rotterdam, from <italic>Salix</italic> sp., 1982, J. de Gruyter,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=101202&link_type=cbs">CBS 101202</ext-link> = PD
82/942.</p><p><italic><bold>Boeremia exigua</bold></italic> var. <italic><bold>pseudolilacis</bold></italic>Aveskamp, Gruyter & Verkley, <bold>var. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515631&link_type=mb">MB515631</ext-link>.
<xref ref-type="fig" rid="fig10">Fig. 10</xref>.</p><p>Varietas haec in cultura habitu Phomae exiguae var. exiguae et var.
gilvescentis similis, sed matrix conidiorum roseo-bubalina et citius
crescens.</p><p><italic>Etymology</italic>: Refers to the former placement in and close resemblance
to <italic>Ph. exigua</italic> var. <italic>lilacis</italic>.</p><p><italic>Colonies</italic> on OA 70–75 mm diam after 7 d, margin regular.
Immersed mycelium black to greenish olivaceous, sparsely visible due to
coverage by a mat of mouse-grey woolly to compact aerial mycelium; reverse
mouse-grey to olivaceous. Colonies on MEA 70–75 mm diam. after 7 d,
margin regular. Immersed mycelium completely covered by a compact aerial mat,
which is smoke-grey with some mouse-grey to white zones; reverse black.
Colonies on CHA slower growing, 70–80 mm diam after 7 d, margin regular,
appearance similar as on MEA. Application of NaOH did not have any effect.
Pycnidial and conidial shapes and sizes fit within the <italic>B. exigua</italic>species concept. Conidial matrix rosy-buff.</p><p><italic>Specimen examined</italic>: <bold>The Netherlands</bold>, near Boskoop, from
<italic>Syringa vulgaris</italic>, 1994, J. de Gruyter, <bold>holotype</bold> CBS H-20371,
culture ex-holotype <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=101207&link_type=cbs">CBS
101207</ext-link> = PD 94/614.</p><p><italic>Notes</italic>: This novel variety of <italic>B. exigua</italic>, distinguished
from other <italic>B. exigua</italic> varieties on basis of DAF analysis
(<xref ref-type="bibr" rid="ref20">Aveskamp <italic>et al.</italic>2009b</xref>) and AFLP (<xref ref-type="bibr" rid="ref8">Abeln <italic>et
al.</italic> 2002</xref>), is closely related to <italic>B. exigua</italic> var.
<italic>exigua</italic> and <italic>B. exigua</italic> var. <italic>gilvescens</italic>. Upon
collection, the strain representing <italic>B. exigua</italic> var.
<italic>pseudolilacis</italic> has probably erroneously been identified as var.
<italic>lilacis</italic> due to its host association.</p><p><italic><bold>Boeremia exigua</bold></italic> var. <italic><bold>viburni</bold></italic> (Roum. ex.
Sacc.) Aveskamp, Gruyter & Verkley, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515633&link_type=mb">MB515633</ext-link>.</p><p><italic>Basionym</italic>: <italic>Ascochyta viburni</italic> Roum. ex Sacc., Syll. Fung.
3: 387. 1884.</p><p><list list-type="simple"><list-item><p>≡ <italic>Phoma exigua</italic> var. <italic>viburni</italic> (Roum. ex. Sacc.)
Boerema apud De Gruyter & P. Scheer, J. Phytopathol. 146: 414. 1998.</p></list-item></list></p><p><italic>Specimens examined</italic>: <bold>The Netherlands</bold>, Boskoop, from
<italic>Viburnum opulus</italic>, 1984, G.H. Boerema,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=100354&link_type=cbs">CBS 100354</ext-link> = PD
83/448; from <italic>Lonicera</italic> sp., 1993, J. de Gruyter,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=101211&link_type=cbs">CBS 101211</ext-link> = PD
93/838.</p><p><italic><bold>Boeremia foveata</bold></italic> (Foister) Aveskamp, Gruyter & Verkley,
<bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515653&link_type=mb">MB515653</ext-link>.</p><p><italic>Basionym</italic>: <italic>Phoma foveata</italic> Foister, Trans. & Proc. Bot.
Soc. Edinburgh 33: 66. 1940.</p><p><italic>Specimens examined</italic>: <bold>Bulgaria</bold>, from a tuber of <italic>Solanum
tuberosum</italic>, 1994, J. de Gruyter,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=109176&link_type=cbs">CBS 109176</ext-link> = CECT
2828 = PD 94/1394. <bold>U.K.</bold>, from a tuber of <italic>Solanum tuberosum</italic>,
Mar. 1937, C.E. Foister, ex-isotype culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=200.37&link_type=cbs">CBS 200.37</ext-link>;
Northern Ireland, Belfast, from a tuber of <italic>Solanum tuberosum</italic>, 1966,
C. Logan, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=341.67&link_type=cbs">CBS
341.67</ext-link> = CECT 20055 = IMI 331912.</p><p><italic><bold>Boeremia hedericola</bold></italic> (Durieu & Mont.) Aveskamp, Gruyter
& Verkley, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515634&link_type=mb">MB515634</ext-link>.</p><p><italic>Basionym</italic>: <italic>Phyllosticta hedericola</italic> Durieu & Mont.,
Flore d'Algérie Cryptog. 1: 611. 1849.</p><p><list list-type="simple"><list-item><p>≡ <italic>Phoma hedericola</italic> (Durieu & Mont.) Boerema, Trans.
Brit. Mycol. Soc. 67: 295. 1976.</p></list-item></list></p><p><italic>Specimens examined</italic>: <bold>The Netherlands</bold>, Meppel, from a leaf of
<italic>Hedera helix</italic>, 1970,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=366.91&link_type=cbs">CBS 366.91</ext-link> = PD
70/811; from <italic>Hedera helix</italic>, 1987, J. de Gruyter,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=367.91&link_type=cbs">CBS 367.91</ext-link> = PD
87/229.</p><p><italic>Note</italic>: Strain <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=367.91&link_type=cbs">CBS
367.91</ext-link> is sterile.</p><p><italic><bold>Boeremia lycopersici</bold></italic> (Cooke) Aveskamp, Gruyter &
Verkley, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515635&link_type=mb">MB515635</ext-link>.</p><p><italic>Basionym</italic>: <italic>Phoma lycopersici</italic> Cooke, Grevelia 13: 94. 1885.
Teleomorph: <italic>Didymella lycopersici</italic> Kleb., Z. Pflanzenkrankh. 31: 9.
1921.</p><p><italic>Specimens examined</italic>: <bold>The Netherlands</bold>, Heerde, from fruit of
<italic>Lycopersicon esculentum</italic>, Aug. 1967, G.H. Boerema,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=378.67&link_type=cbs">CBS 378.67</ext-link> = PD
76/276; from <italic>Lycopersicon esculentum</italic>, 1984, J. de Gruyter,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=109172&link_type=cbs">CBS 109172</ext-link> = PD
84/143.</p><p><italic><bold>Boeremia noackiana</bold></italic> (Allesch.) Aveskamp, Gruyter &
Verkley, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515636&link_type=mb">MB515636</ext-link>.</p><p><italic>Basionym</italic>: <italic>Phyllosticta noackiana</italic> Allesch., Bol. Inst.
Agron. Campinas 9: 85. 1898.</p><p><list list-type="simple"><list-item><p>≡ <italic>Phoma exigua</italic> var. <italic>noackiana</italic> (Allesch.) Aa,
Boerema & Gruyter, Persoonia 17: 450. 2000.</p></list-item></list></p><p>For a complete description see Van der Aa <italic>et al.</italic>(<xref ref-type="bibr" rid="ref7">2000</xref>).</p><p><italic>Specimens examined</italic>: <bold>Colombia</bold>, from <italic>Phaseolus
vulgaris</italic>, 1979, J. de Gruyter,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=101203&link_type=cbs">CBS 101203</ext-link> = PD
79/1114. <bold>Guatemala</bold>, from <italic>Phaseolus vulgaris</italic>, 1987, IPO
Wageningen, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=100353&link_type=cbs">CBS
100353</ext-link> = PD 87/718.</p><p><italic>Notes</italic>: <italic>Boeremia noackiana</italic> is genetically a sister species
to <italic>B. diversispora</italic>, and was also noted by Boerema <italic>et al</italic>.
(<xref ref-type="bibr" rid="ref55">2004</xref>) as “the American
cousin”. Just like <italic>B. diversispora</italic>, the present species is
known from beans, although the main host appears to be <italic>Phaseolus
vulgaris.</italic> The two species have many characters in common with <italic>B.
exigua</italic> (<xref ref-type="bibr" rid="ref7">Van der Aa <italic>et al.</italic>2000</xref>, <xref ref-type="bibr" rid="ref55">Boerema <italic>et al.</italic>2004</xref>) and with each other, but are distinguished based on enzyme
analysis (<xref ref-type="bibr" rid="ref154">Obando-Rojas, 1989</xref>)
and molecular fingerprinting methods such as AFLP
(<xref ref-type="bibr" rid="ref8">Abeln <italic>et al.</italic> 2002</xref>)
and DAF (<xref ref-type="bibr" rid="ref20">Aveskamp <italic>et al.</italic>2009b</xref>). Additionally, <italic>B. noackiana</italic> is characterised by a
relative fast growth rate on MEA: (6–)6.5–7.5 mm diam after 7 d,
and is further distinguished from <italic>B. diversispora</italic> by its relatively
uniform conidia. Due to the relatively large genetical distance to the <italic>B.
exigua</italic> complex, this taxon is elevated to species level.</p><p><italic><bold>Boeremia sambuci-nigrae</bold></italic> (Sacc.) Aveskamp, Gruyter &
Verkley, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515637&link_type=mb">MB515637</ext-link>.</p><p><italic>Basionym</italic>: <italic>Phoma herbarum</italic> f. <italic>sambuci-nigrae</italic>Sacc., Syll. Fung. 3: 133. 1884.</p><p><list list-type="simple"><list-item><p>≡ <italic>Phoma exigua</italic> var. <italic>sambuci-nigrae</italic> (Sacc.) Boerema
& Höweler, Persoonia 5(1): 26. 1967.</p></list-item><list-item><p>≡ <italic>Phoma sambuci-nigrae</italic> (Sacc.) E. Monte, Bridge & B.
Sutton, Mycopathologia 115: 102. 1991.</p></list-item></list></p><p><italic>Specimens examined</italic>: <bold>The Netherlands</bold>, Wageningen, from a
leaf of <italic>Sambucus nigra</italic>, 1967, <bold>lectotype</bold> CBS H-16314,
ex-lectotype culture <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=629.68&link_type=cbs">CBS
629.68</ext-link> = CECT 20048 = IMI 331913 = PD 67/753; Baarn,
Maarschalksbos, from a leaf of <italic>Sambucus nigra</italic>, Nov. 1967, H.A. van
der Aa, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=104.68&link_type=cbs">CBS 104.68</ext-link>=
CECT 20010; from <italic>Sambucus nigra</italic>, 1975, G.H. Boerema,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=109170&link_type=cbs">CBS 109170</ext-link> = PD
75/796.</p><p><italic><bold>Boeremia strasseri</bold></italic> (Moesz) Aveskamp, Gruyter & Verkley,
<bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515638&link_type=mb">MB515638</ext-link>.</p><p><italic>Basionym</italic>: <italic>Phoma strasseri</italic> Moesz, <italic>Bot.
Közlem.</italic> 22: 45. 1924. nom. nov. pro <italic>Phoma menthae</italic> Strasser,
Verh. zool. Bot. Ges. Wien 60: 317. 1910 [non <italic>Phoma menthae</italic> Roum.
(date of publication unknown)].</p><p><italic>Specimens examined</italic>: <bold>The Netherlands</bold>, Arnhem, from a stem of
<italic>Mentha</italic> sp., 1973, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=126.93&link_type=cbs">CBS
126.93</ext-link> = PD 73/642. <bold>U.S.A.</bold>, Oregon, from <italic>Mentha
piperita</italic>, 1970, H.A. van der Aa,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=261.92&link_type=cbs">CBS 261.92</ext-link> = ATCC
244146 = PD 92/318.</p><p><italic>Note</italic>: As the older name <italic>Ph. menthae</italic> is illegitimate, the
epithet “<italic>strasseri</italic>” prevails.</p><p><italic><bold>Boeremia telephii</bold></italic> (Vestergr.) Aveskamp, Gruyter &
Verkley, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515639&link_type=mb">MB515639</ext-link>.</p><p><italic>Basionym</italic>: <italic>Ascochyta telephii</italic> Vestergr<italic>.,</italic>Öfvers. Förh. Kongl. Svenska Vetenska.-Akad. 54: 41. 1897.</p><p><list list-type="simple"><list-item><p>≡ <italic>Phoma telephii</italic> (Vestergr.) Kesteren, Netherlands J. Pl.
Pathol. 78: 117.1972.</p></list-item></list></p><p><italic>Specimens examined</italic>: <bold>The Netherlands</bold>, Utrecht, from a stem
of <italic>Sedum telephium</italic>, 1971, G.H. Boerema,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=760.73&link_type=cbs">CBS 760.73</ext-link> = PD
71/1616; from <italic>Sedum spectabile</italic>, 1975, G.H. Boerema,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=109175&link_type=cbs">CBS 109175</ext-link> = PD
79/524.</p></sec><sec><title>Group O:</title><p>Three species are clustered in group O, which all were accommodated in the
Boeremaean section <italic>Phoma</italic>. These species, <italic>Ph. multirostrata, Ph.
pereupyrena</italic> and <italic>Ph. insulana</italic> are characterised by the production
of small (5–15 μm diam), unicellular chlamydospores, comparable to
those formed by some species in group K. The absence of septate conidia and a
thin pycnidial wall are further characters of the species accommodated in
group O.</p><p>The strains accommodated in <italic>Ph. multirostrata</italic> reveal a high
variation in spore size. Boerema <italic>et al.</italic> (1986) introduced three
varieties within this species, but based on morphological observations and DNA
sequence analyses, these varieties were not recognised by later researchers
and thus the varieties were merged again
(<xref ref-type="bibr" rid="ref140">Morgan-Jones 1988</xref>,
<xref ref-type="bibr" rid="ref19">Aveskamp <italic>et al.</italic>2009a</xref>).</p><p><italic><bold>Phoma insulana</bold></italic> (Mont.) Boerema & Malathr., in Boerema,
Verslagen Meded. Plziektenk. Dienst Wageningen 158 (Jaarboek 1981): 28.
1982.</p><p><italic>Basionym</italic>: <italic>Phyllosticta insulana</italic> Mont., Ann. Sci. Nat.
Bot. IV 5: 343.1856.</p><p><italic>Specimen examined</italic>: <bold>Greece</bold>, from the berries of <italic>Olea
europaea</italic>, 1980, G.H. Boerema,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=252.92&link_type=cbs">CBS 252.92</ext-link> = PD
80/1144.</p><p><italic><bold>Phoma multirostrata</bold></italic> (P.N. Mathur, S.K. Menon & Thirum.)
Dorenb. & Boerema, Mycopathol. Mycol. Appl. 50(3): 256. 1973, emend.
Aveskamp <italic>et al.</italic> Mycologia 101: 375. 2009.</p><p><italic>Basionym</italic>: <italic>Sphaeronaema multirostratum</italic> P.N. Mathur, S.K.
Menon & Thirum., Sydowia 13: 146. 1959. (as
“<italic>Sphaeronema</italic>”).</p><p><italic>Specimens examined</italic>: <bold>India</bold>, Maharashtra, Poona, Talegaon,
from poultry farm soil, Mar. 1959, M.J. Thirumalachar, <bold>isotype</bold> CBS
H-7616, culture <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=274.60&link_type=cbs">CBS
274.60</ext-link> = IMI 081598; Maharashtra, Poona, Talegaon, from soil, Mar.
1959, M.J. Thirumalachar, CBS H-16499, culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=368.65&link_type=cbs">CBS 368.65</ext-link> = PD
92/1757. <bold>The Netherlands</bold>, Hoorn, greenhouse, from the stem of
<italic>Cucumis sativus</italic>, Aug. 1967, G.H. Boerema, CBS H-16502, culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=110.79&link_type=cbs">CBS 110.79</ext-link> = PD
65/8875. Unknown origin, from <italic>Cucumis sativus</italic>, 1983, PD 83/48.</p><p><italic><bold>Phoma pereupyrena</bold></italic> Gruyter, Noordel. & Boerema,
Persoonia 15(3): 390. 1993.</p><p><italic>Specimen examined</italic>: <bold>India</bold>, from a leaf of <italic>Coffea
arabica,</italic> 1976, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=267.92&link_type=cbs">CBS
267.92</ext-link> = PD 76/1014.</p></sec><sec><title>Group P:</title><p>This well-supported clade (BPP = 1.00, RBS = 97 %) comprises <italic>Ph.
dictamnicola</italic> and <italic>Ph. sylvatica</italic>, which are both associated with
the section <italic>Sclerophomella</italic> (Boerema <italic>et al.</italic> 1998). In
addition, both varieties of <italic>Ph. poolensis</italic> are recovered here. As in
the <italic>Sclerophomella</italic> species, an ostiole is commonly absent in <italic>Ph.
poolensis</italic> var. <italic>poolensis</italic>, a character which supports the
sequence data found in the present study. In contrast, the second variety of
this species, <italic>Ph. poolensis</italic> var. <italic>verbascicola</italic>, always
produces ostiolate pycnidia (<xref ref-type="bibr" rid="ref97">De Gruyter
<italic>et al.</italic> 1993</xref>). Both <italic>Ph. poolensis</italic> varieties can
further be differentiated on the basis of the β-tubulin sequence, and are
morphologically distinguishable in the colour of the conidial matrix. The
conidia of the type variety are on average somewhat smaller, measuring
<italic>ca</italic>. 3.5–5 × 1.5–2 μm, than those of var.
<italic>verbasicola</italic>, which measure 3.5–5.5 × 1.5–2.5 μm.
Both varieties are known from plant hosts belonging to the
<italic>Scophulariaceae</italic>, but whereas var. <italic>poolensis</italic> is recorded as
causal agent of leaf spots and basal stem rot in snapdragon (<italic>Antirrhinum
majus</italic>), var. <italic>verbascicola</italic> is only known as saprobe of
<italic>Verbascum</italic> spp., although inoculation trials indicated that it may
also have a role as pathogen (<xref ref-type="bibr" rid="ref55">Boerema
<italic>et al.</italic> 2004</xref>). Given all these differences, it is
considered to be justified to erect a separate species for <italic>Ph.
poolensis</italic> var. <italic>verbascicola</italic> as <italic>Ph.
novae-verbascicola</italic>.</p><p>Although none of the species in this group has been confirmed to have a
teleomorph (Boerema <italic>et al.</italic> 1998), it has been suggested that
<italic>Didymella winteriana</italic> is the teleomorph of <italic>Phoma sylvatica</italic>(<xref ref-type="bibr" rid="ref144">Munk 1957</xref>). Given the
topology of the tree, this association with a <italic>Didymella</italic> species is
plausible, although a sexual structure was not observed in the present study,
nor in the previous studies of Boerema & De Gruyter
(<xref ref-type="bibr" rid="ref40">1998</xref>).</p><p><italic><bold>Phoma dictamnicola</bold></italic> Boerema, Gruyter & Noordel.,
Persoonia 15(1): 90. 1992.</p><p><italic>Specimen examined</italic>: <bold>The Netherlands</bold>, Arnhem, from a stem of
<italic>Dictamnus albus</italic>, 1974, J. de Gruyter,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=507.91&link_type=cbs">CBS 507.91</ext-link> = PD
74/148.</p><p><italic><bold>Phoma novae-verbascicola</bold></italic> Aveskamp, Gruyter & Verkley,
<bold>nom. nov.</bold> pro <italic>Phyllosticta verbascicola</italic> Ellis & Kellerm.
MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515640&link_type=mb">MB515640</ext-link>.</p><p><italic>Basionym</italic>: <italic>Phyllosticta verbascicola</italic> Ellis & Kellerm.,
Bull. Torrey Bot. Club 11: 115. 1884.</p><p><list list-type="simple"><list-item><p>≡ <italic>Phoma poolensis</italic> var. <italic>verbascicola</italic> (Ellis &
Kellerm.) Aa & Boerema, in De Gruyter, Noordeloos & Boerema, Persoonia
15(3): 385. 1993. Not <italic>Phoma verbascicola</italic> (Schwein.) Cooke, in
Ravenel. 1878.</p></list-item></list></p><p><italic>Etymology</italic>: The epithet refers to the host plant,
<italic>Verbascum</italic> spp.</p><p>For a full description see De Gruyter <italic>et al.</italic>(<xref ref-type="bibr" rid="ref97">1993</xref>).</p><p><italic>Specimens examined</italic>: <bold>The Netherlands</bold>, Zeist, Abburg nursery,
<bold>holotype</bold> L 9893.00.134; Haarlem, from dead stem material of
<italic>Verbascum densiflorum</italic>, 1992, J. de Gruyter,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=127.93&link_type=cbs">CBS 127.93</ext-link> = PD
92/347; from stem of <italic>Verbascum</italic> sp., 1974, G.H. Boerema,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=114.93&link_type=cbs">CBS 114.93</ext-link> = PD
74/228.</p><p><italic>Notes</italic>: This species is distinguishable from <italic>Ph. poolensis</italic>by to the presence of 1–2(–5) ostioles, the colourless to whitish
matrix and the smaller conidia. On MEA, the aerial mycelium is more compact or
woolly than that of <italic>Ph. poolensis</italic>.</p><p>The variety epithet could not be elevated to species level, as <italic>Phoma
verbascicola</italic> is already occupied. This basionym, however, probably refers
to immature pseudothecia of a <italic>Pleospora</italic> species
(<xref ref-type="bibr" rid="ref53">Boerema <italic>et al.</italic>1996</xref>). Therefore, a new name is proposed here for the present
species.</p><p><italic><bold>Phoma poolensis</bold></italic> Taubenh., Dis. Greenhouse Crops 203.
1919.</p><p><italic>Specimens examined</italic>: <bold>Denmark</bold>, from a stem of <italic>Antirrhinum
majus</italic>, July 1938, P. Neergaard,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=253.38&link_type=cbs">CBS 253.38</ext-link>. <bold>The
Netherlands</bold>, Wageningen, from a stem of <italic>Scrophularia nodosa</italic>,
1974, G.H. Boerema, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=115.93&link_type=cbs">CBS
115.93</ext-link> = PD 74/206; Bennekom, from a stem of <italic>Antirrhinum
majus</italic>, 1973, G.H. Boerema,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=116.93&link_type=cbs">CBS 116.93</ext-link>= PD
71/884. Unknown origin and substrate, 1920, E.M. Smiley,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=113.20&link_type=cbs">CBS 113.20</ext-link> = PD
92/774.</p><p><italic><bold>Phoma sylvatica</bold></italic> Sacc. Michelia 2(2): 337. 1881.</p><p><italic>Specimens examined</italic>: <bold>The Netherlands</bold>, Wageningen, from
<italic>Melampyrum pratense</italic>, 1983, J. de Gruyter,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=135.93&link_type=cbs">CBS 135.93</ext-link> = PD
83/87; Wageningen, from a stem of <italic>Melampyrum pratense</italic>, 1993, J. de
Gruyter, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=874.97&link_type=cbs">CBS 874.97</ext-link>= PD 93/764.</p></sec><sec><title>Group Q:</title><p>The <italic>Phoma</italic> species embedded in this group, <italic>Ph.
commelinicicola</italic> and <italic>Ph. eupatorii</italic> are morphologically distinct,
hence their accommodation in the sections <italic>Phoma</italic> and
<italic>Macrospora</italic> respectively. The accommodation of <italic>Chaetasbolisia
erysiphoides</italic> in this clade, the type of its genus, is unexpected.
Attempted morphological studies revealed that this strain was sterile, and
therefore recombination of the species could not be supported by morphological
data. The descriptions provided in literature
(<xref ref-type="bibr" rid="ref219">Sutton 1980</xref>, Patel <italic>et
al.</italic> 1997, <xref ref-type="bibr" rid="ref181">Reynolds 1999</xref>)
suggest, however, that this genus could very well represent a group of setose
<italic>Phoma</italic> species, although this cannot be resolved due to a lack of
isolates. The presence of setae is not recorded in other species in group Q,
and moreover, is within the <italic>Didymellaceae</italic> only known from
<italic>Peyronellaea gardeniae</italic> (Group K), and from pycnidia in some older
cultures from <italic>Epicoccum sorghi</italic> (Group M), <italic>Peyronellaea
glomerata</italic> (Group K) and <italic>Phoma herbarum</italic>(<xref ref-type="bibr" rid="ref55">Boerema <italic>et al</italic>.
2004</xref>). The topology and the clustering of these species cannot be
further explained by the morphology or ecology, nor by their geographical
distribution.</p><p><italic><bold>Phoma commelinicola</bold></italic> (E. Young) Gruyter, Persoonia 18(1):
93. 2002.</p><p><italic>Basionym</italic>: <italic>Phyllosticta commelinicola</italic> E. Young, Mycologia
7: 144. 1915.</p><p><italic>Specimen examined</italic>: <bold>New Zealand</bold>, South Auckland, Alfriston,
from <italic>Tradescantia</italic> sp., 1997, K. Ramsay,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=100409&link_type=cbs">CBS 100409</ext-link>.</p><p><italic><bold>Phoma eupatorii</bold></italic> Died., Ann. Mycol. 10(5): 447. 1912.</p><p><italic>Specimen examined</italic>: <bold>The Netherlands</bold>, Arnhem, from
<italic>Eupatorium cannabinum</italic>, 1977, G.H. Boerema,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123.93&link_type=cbs">CBS 123.93</ext-link> = PD
77/1148.</p></sec><sec><title>Group R:</title><p>This group comprises five species that previously were accommodated in the
sections <italic>Phoma, Peyronellaea</italic> and <italic>Phyllostictoides</italic>. As the
name of <italic>Ph. tropica</italic> already suggests, it concerns a thermotolerant
species, which is mainly found in European greenhouses on a wide range of
hosts, but which probably has a tropical origin
(<xref ref-type="bibr" rid="ref192">Schneider & Boerema
1975</xref>), as do most other species found in the present clade. The
sole host of <italic>Ph. costarricensis</italic> is coffee bean (<italic>Coffea
arabica</italic>), while <italic>Ph. piperis</italic> is associated with Indian Long
Pepper (<italic>Piper longus</italic>), and the novel species <italic>Ph. minor</italic> has
been isolated twice from clove (<italic>Syzygium aromaticum</italic>) in Indonesia. In
addition, <italic>Ph. labilis</italic> is a warmth-preferring plurivorous species that
has been isolated in European greenhouses and from nature in the Middle East,
Turkey and Indonesia (<xref ref-type="bibr" rid="ref55">Boerema <italic>et
al.</italic> 2004</xref>). <italic>Phoma zantedeschiae</italic> is widespread
throughout the Western Hemisphere, but always in association with arum or
calla (<italic>Zantedeschia</italic> sp.), a genus that is indigenous in southern
Africa (<xref ref-type="bibr" rid="ref39">Boerema & Hamers
1990</xref>). Thus far, however, no data of temperature-growth studies
are available for these species except for <italic>Ph. tropica</italic>. Several other
thermotolerant species, such as <italic>Ph. calidophila, Ph. calorpreferens</italic>and <italic>Ph. multirostrata</italic>, are, however, not accommodated in this group.
These three species are soil-borne, in contrast to <italic>Ph. tropica</italic> and
<italic>Ph. costarricensis</italic>, which are associated with leaf-spots.</p><p><italic>Phoma tropica</italic> and <italic>Ph. costarricensis</italic> are both closely
related, and colony characters are highly similar. However, the strains
available revealed a significant difference in conidial and pycnidial sizes,
consistent with the data obtained in previous studies
(<xref ref-type="bibr" rid="ref192">Schneider & Boerema
1975</xref>, <xref ref-type="bibr" rid="ref96">De Gruyter &
Noordeloos 1992</xref>).</p><p><italic><bold>Phoma costarricencis</bold></italic> Echandi, Rev. Biol. Trop. 5: 83.
1957.</p><p><italic>Specimens examined</italic>: <bold>Nicaragua</bold>, from a twig of
<italic>Coffea</italic> sp., 1991, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=506.91&link_type=cbs">CBS
506.91</ext-link> = PD 91/876 = IMI 215229. Unknown origin, from <italic>Coffea
arabica</italic>, 1979, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=497.91&link_type=cbs">CBS
497.91</ext-link> = PD 79/209.</p><p><italic>Notes</italic>: Strain <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=497.91&link_type=cbs">CBS
497.91</ext-link> was initially identified as <italic>Ph. tropica</italic>. The close
phylogenetic association between this species and <italic>Ph. costarricensis</italic>concurs with their overlapping morphological characters (see
<xref ref-type="bibr" rid="ref192">Schneider & Boerema 1975</xref>,
<xref ref-type="bibr" rid="ref96">De Gruyter & Noordeloos
1992</xref>).</p><p><italic><bold>Phoma labilis</bold></italic> Sacc., Michelia 2(7): 341. 1881.</p><p><italic>Specimens examined</italic>: <bold>Israel</bold>, from a stem of <italic>Rosa</italic>sp., 1970, G.H. Boerema, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=479.93&link_type=cbs">CBS
479.93</ext-link> = PD 70/93. <bold>The Netherlands</bold>, Barendrecht, from a
stem of <italic>Lycopersicon esculentum</italic>, 1987, J. de Gruyter,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=124.93&link_type=cbs">CBS 124.93</ext-link> = PD
87/269.</p><p><italic><bold>Phoma minor</bold></italic> Aveskamp, Gruyter & Verkley, <bold>sp.
nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515641&link_type=mb">MB515641</ext-link>.
<xref ref-type="fig" rid="fig11">Fig. 11</xref>.</p><p>Conidia ellipsoidea, ovoidea vel leniter allantoidea, glabra, hyalina,
continua, (3–)3.5–4.5(–5) × 1.8–2.5(–3)
μm, (0–)1–3(–4) guttulis minutis praedita. Matrix
conidiorum alba.</p><p><italic>Etymology</italic>: Epithet derived from the small-sized conidia.</p><p><italic>Conidiomata</italic> pycnidial, solitary, (sub-)globose to broadly
ellipsoidal, glabrous or with some hyphal outgrows, on the agar surface and
immersed, (125–)150–280(–330) ×
(125–)150–220(–245) μm. <italic>Ostioles</italic> (1–5),
slightly papillate or non-papillate. <italic>Pycnidial wall</italic>pseudoparenchymatous, composed of oblong to isodiametric cells, outer cell
layer pigmented, 2–4 layers, 8–15 μm thick. <italic>Conidiogenous
cells</italic> phialidic, hyaline, simple, smooth, flask-shaped or somewhat
isodiametric, <italic>ca</italic>. 4–5.5(–6.2) ×
3–4.5(–4.7) μm. <italic>Conidia</italic> ellipsoidal to ovoid or
slightly allantoid, thin-walled, smooth, hyaline, aseptate
(3–)3.5–4.5(–5) × 1.8–2.5(–3) μm, with
(0–)1–3(–4) minute guttules. Conidial matrix white.</p><p><italic>Culture characteristics</italic>: Colonies on OA
(44–)45–50(–54) mm diam after 7 d, margin regular. Aerial
mycelium flat, grey, but locally well-developed in densely floccose white
tufts. Immersed mycelium olivaceous with rosy-buff tinges near the colony
margin; reverse concolourous. Colonies on MEA 46-48 mm diam after 7 d, margin
regular. Immersed mycelium hyaline, with abundant semi-immersed pycnidia, but
almost completely covered by an aerial mycelial mat. Aerial mycelium
pluriform, with a compact white mat and some felty glaucous grey or dull green
zones, near colony margin white; reverse black to grey-olivaceous. Colonies on
CHA 50–54 mm diam. after 7 d, margin regular. Aerial mycelium similar as
on MEA, although the felty white and glaucous grey zones are less abundant;
reverse slate blue to leaden-black. Application of NaOH results in a greenish
yellow discolouration of the agar, best to be observed on OA medium.</p><p><italic>Specimens examined</italic>: <bold>Indonesia</bold>, Sumatra, from <italic>Syzygium
aromaticum</italic>, Apr. 1982, R. Kasim, <bold>holotype designated here</bold> CBS
H-20236, ex-holotype culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=325.82&link_type=cbs">CBS 325.82</ext-link>;
Lampung, from <italic>Syzygium aromaticum</italic>, Dec. 1982, H. Vermeulen,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=315.83&link_type=cbs">CBS 315.83</ext-link>.</p><p><italic>Notes</italic>: As for <italic>Ph. eucalyptica</italic>, this species has been
recorded in association with clove trees (<italic>Syzygium aromaticum</italic>,
<xref ref-type="bibr" rid="ref55">Boerema <italic>et al.</italic> 2004</xref>).
Both species, although genetically distinct, have many characters in common,
notably the colony characters on OA, the high variation in ostiole number and
a similar reaction to application of NaOH. Although <italic>Phoma minor</italic>produces relatively small conidia, the conidia of <italic>Ph. eucalyptica</italic> are
even smaller, measuring only 2–4 × 1–2 μm
(<xref ref-type="bibr" rid="ref96">De Gruyter & Noordeloos
1992</xref>).</p><p><italic><bold>Phoma piperis</bold></italic> (Tassi) Aa & Boerema, Persoonia 15(3):
398. 1993.</p><p><italic>Basionym</italic>: <italic>Phyllosticta piperis</italic> Tassi, Boll. Reale Orto
Bot. Siena 3(2): 28. 1900.</p><p><italic>Specimens examined</italic>: <bold>The Netherlands</bold>, Tiel, from a leaf of
<italic>Peperomia pereskiifolia</italic>, 1988, J. de Gruyter,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=268.93&link_type=cbs">CBS 268.93</ext-link> =
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=108.93&link_type=cbs">CBS 108.93</ext-link> = PD
88/720; Tiel, from <italic>Peperomia</italic> sp., 1990, J. de Gruyter, PD 90/2011</p><p><italic><bold>Phoma tropica</bold></italic> R. Schneid. & Boerema, Phytopathol. Z. 83
(4): 361. 1975.</p><p><italic>Specimen examined</italic>: <bold>Germany</bold>, Horrheim, from <italic>Saintpaulia
ionantha</italic>, 1973, R. Schneider, <bold>isotype</bold> CBS H-7629, ex-isotype
culture <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=436.75&link_type=cbs">CBS
436.75</ext-link>.</p><p><italic><bold>Phoma zantedeschiae</bold></italic> Dippen., S. African J. Sci. 28: 284.
1931.</p><p><italic>Specimen examined</italic>: <bold>The Netherlands</bold>, from a bulb of
<italic>Zantedeschiae</italic> sp., 1969, G.H. Boerema,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=131.93&link_type=cbs">CBS 131.93</ext-link> = PD
69/140.</p></sec><sec><title>Group S – Stagonosporopsis:</title><p>This large group (BPP = 1.00, RBS = 55 %) comprises mainly species with
<italic>Stagonosporopsis</italic> synanamorphs. In the Boeremaean classification
system, these species were embedded in <italic>Phoma</italic> section
<italic>Heterospora</italic> (<xref ref-type="bibr" rid="ref54">Boerema <italic>et
al.</italic> 1997</xref>). As with the other sections, this group also
appeared to be artificial. Based on LSU and SSU sequences, the type species of
the section <italic>Heterospora, Ph. heteromorphospora</italic>, clusters outside the
<italic>Didymellaceae</italic> (<xref ref-type="bibr" rid="ref100">De Gruyter <italic>et
al</italic>. 2009</xref>), as do <italic>Ph. samarorum</italic> and <italic>Ph.
dimorphospora</italic>. Three species, <italic>Ph. clematidina, Ph. glaucii</italic> and
<italic>Ph. aquilegiicola</italic> form a separate clade (Group C) within the
<italic>Didymellaceae</italic>, and are treated above. Also <italic>Ph. nigripycnidia</italic>and <italic>Ph. narcissi</italic> are not accommodated here.</p><p><fig position="float" id="fig11"><label>Fig. 11.</label><caption><p><italic>Phoma minor</italic> (<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=325.82&link_type=cbs">CBS
325.82</ext-link>). A–C. Fourteen-day-old colonies on OA (A), MEA (B)
and CHA (C). D–E. Pycnidia. F. Section of the pycnidial wall. G.
Conidia. Scale bars: D = 200 μm; E = 100 μm; F–G = 10 μm.</p></caption><graphic xlink:href="1fig11"></graphic></fig></p><p>In contrast to the <italic>Heterospora</italic> species that are absent in this
clade, several current <italic>Phoma</italic> taxa recovered here have been associated
with the section <italic>Phyllostictoides</italic>, such as <italic>Ph. artemisiicola, Ph.
caricae-papayae, Ph. cucurbitacearum, Ph. heliopsidis, Ph. rudbeckiae</italic>,
and the quarantine-organisms <italic>Ph. ligulicola</italic> var. <italic>ligulicola</italic>and var. <italic>inoxydabilis</italic> (<xref ref-type="bibr" rid="ref94">De Gruyter
2002</xref>). These are all included in subclade S1 (BPP = 0.93, RBS = 73
%). These species do produce a percentage of multicellular conidia in culture
that are often considerably larger than the regular aseptate ones. However,
Boerema <italic>et al.</italic> (<xref ref-type="bibr" rid="ref54">1997</xref>)
decided to exclude the <italic>Ph. ligulicola</italic> varieties and <italic>Ph.
cucurbitacearum</italic> from section <italic>Heterospora</italic>, as the sizes of the
<italic>Stagonosporopsis</italic>-like conidia do not always exceed that of the
aseptate conidia in these species. A sister clade to subclade S1 is S2, which
hosts the potato pathogens <italic>Ph. andigena</italic> and <italic>Ph.
crystalliniformis</italic> – formerly known as <italic>Ph. andina</italic> var.
<italic>crystalliniformis</italic>. Both species originate from the Andes region, and
are regarded as serious quarantine pathogens in large parts of the world
(<xref ref-type="bibr" rid="ref208">Smith <italic>et al</italic>.
1992</xref>).</p><p>In addition, three other subclades can be recognised in this clade. One
(S3) comprises the species <italic>Ph. schneiderae</italic> and <italic>Ph.
subboltshauserii</italic> (both of the section <italic>Heterospora</italic>) as well as
<italic>Ph. astragali</italic>. This species is known as a pathogen of
<italic>Astragalus</italic> spp., and is characterised by a high percentage of
“distorted” conidia, but thus far, no records have been made of a
<italic>Stagonosporopsis</italic>-like synanamorph. Whereas records of <italic>Ph.
astragali</italic> and <italic>Ph. schneiderae</italic> are mainly limited to the American
continent, <italic>Ph.</italic> <italic>subboltshauseri</italic> appears to occur worldwide on
<italic>Fabaceae</italic>. However, Boerema <italic>et al.</italic>(<xref ref-type="bibr" rid="ref55">2004</xref>) suggested that the
original host of this species may have been <italic>Phaseolus</italic>, which is
native to the Americas.</p><p><fig position="float" id="fig12"><label>Fig. 12.</label><caption><p>Conidial dimorphism in three species of <italic>Stagonosporopsis.</italic> A.
<italic>S. actaeae</italic> (<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=106.96&link_type=cbs">CBS
106.96</ext-link>). B. <italic>S. lupini</italic>(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=101494&link_type=cbs">CBS 101494</ext-link>). C.
<italic>S. cucurbitacearum</italic>(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=109171&link_type=cbs">CBS 109171</ext-link>). Scale
bars: A = 20 μm; B–C = 10 μm.</p></caption><graphic xlink:href="1fig12"></graphic></fig></p><p>A fourth and fifth (S4, S5) subclade in this group comprise species that
are accommodated in section <italic>Phoma</italic>, and therefore lack any further
features than a plain, globose pycnidium and aseptate, hyaline conidia. The
species found here are <italic>Ph. dorenboschiae, Ph. loticola</italic> (both S4),
<italic>Ph. ajacis</italic> and <italic>Ph. valerianellae</italic> (both S5).</p><p>In group S several taxa have been found with a teleomorph in
<italic>Didymella</italic>, such as <italic>Ph. ligulicola</italic> var. <italic>ligulicola</italic>(teleomorph <italic>D. ligulicola</italic> var. <italic>ligulicola</italic>), <italic>Ph.
ligulicola</italic> var. <italic>inoxydabilis</italic> (<italic>D. ligulicola</italic> var.
<italic>inoxydabilis</italic>), and <italic>Ph. cucurbitacearum</italic> (<italic>D.
bryoniae</italic>). Also the teleomorph of <italic>Ph. caricae-papayae</italic> has been
recovered in this study, and found to be a <italic>Didymella</italic>, which is in
line with the other teleomorph observations in this clade. The current
teleomorph state of this species is accommodated in <italic>Mycosphaerella</italic> as
<italic>M. caricae</italic> (<xref ref-type="bibr" rid="ref207">Sivanesan
1990</xref>).</p><p>As the species in the present clade form a well-defined group within the
<italic>Didymellaceae</italic>, the taxa are recombined into the genus
<italic>Stagonosporopsis</italic>. This further implies that the names of the
<italic>Stagonosporopsis</italic> synanamorphs of <italic>Ph. samarorum</italic> and <italic>Ph.
narcissi</italic> (<italic>S. fraxini</italic> and <italic>S. curtisii</italic> respectively)
should no longer be used.</p><p><italic><bold>Stagonosporopsis</bold></italic> Died., Ann. Mycol. 10(2): 142. 1912.
<bold>emend.</bold> Aveskamp, Gruyter & Verkley.
<xref ref-type="fig" rid="fig12">Fig. 12</xref>.</p><p><italic>Conidiomata</italic> pycnidial, globose to subglobose, measuring
70–300 μm diam, on agar surface or immersed, solitary or confluent,
ostiolate or poroid. <italic>Pycnidial wall</italic> pseudoparenchymatous, counting
2–6 cell layers of which the outer 1–3 are brown/olivaceous
pigmented. <italic>Conidiogenous cells</italic> phialidic, hyaline, simple, smooth,
ampulliform or doliiform, <italic>ca</italic>. 4–7.5 × 3–6 μm.
<italic>Conidia</italic> often in two types: majority aseptate, hyaline, ellipsoidal
to subglobose, thin-walled, smooth, measuring (3–)3.5–10 ×
1.5–3(–3.5) μm. Conidia of the second type can be produced both
<italic>in vivo</italic> and <italic>in vitro</italic> in the same pycnidia as the smaller
spores, unicellular or with up to 3 septa, measuring up to 30 × 8 μm.
<italic>Pseudothecia</italic>, if present, occurring only <italic>in vivo</italic>, globose to
subglobose, sometimes with a somewhat conical neck, measuring 90–230
μm diam. <italic>Asci</italic> cylindrical or subclavate, measuring 50–90
× 9–13 μm, always 8-spored, biseriate. <italic>Ascospores</italic>ellipsoid, fusiform or obovoid, measuring 12–18 × 4–7 μm,
uniseptate, guttulate.</p><p><italic><bold>Stagonosporopsis actaeae</bold></italic> (Allesch.) Died., Ann. Mycol. 10:
141. 1912.</p><p><italic>Basionym</italic>: <italic>Actinonema actaeae</italic> Allesch., Ber. bayer. bot.
Ges. 5: 7. 1897.</p><p><list list-type="simple"><list-item><p><italic>= Phoma actaeae</italic> Boerema, Gruyter & Noordeloos, Persoonia
16(3): 347. 1997.</p></list-item></list></p><p><italic>Specimens examined</italic>: <bold>The Netherlands</bold>, Zeist, from a stem of
<italic>Cimicifuga simplex</italic>, 1974, G.H. Boerema,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=105.96&link_type=cbs">CBS 105.96</ext-link> = PD
74/230; Limburg, Schaersbergerbos, from a leaf of <italic>Actaea spicata</italic>,
1994, J. de Gruyter, L 992.167.501, culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=106.96&link_type=cbs">CBS 106.96</ext-link> = PD
94/1318.</p><p><italic>Notes</italic>: In contrast to the earlier description of the
<italic>Phoma</italic> anamorph of this species
(<xref ref-type="bibr" rid="ref54">Boerema <italic>et al.</italic>1997</xref>), the larger conidia regularly produces up to 3-septate
conidia (see <xref ref-type="fig" rid="fig12">Fig. 12A</xref>). In the
study mentioned above and in the present one the same strains were examined
morphologically.</p><p><italic><bold>Stagonosporopsis ajacis</bold></italic> (Thüm.) Aveskamp, Gruyter
& Verkley, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515653&link_type=mb">MB515653</ext-link>.</p><p><italic>Basionym</italic>: <italic>Phyllosticta ajacis</italic> Thüm., apud Bolle
& von Thümen, Boll. Soc. Adriat. Sci. Nat. Trieste 6: 329. 1880.</p><p><list list-type="simple"><list-item><p>= <italic>Phoma ajacis</italic> Aa & Boerema, apud De Gruyter, Noordeloos &
Boerema, Persoonia 15(3): 383. 1993.</p></list-item></list></p><p><italic>Specimens examined</italic>: <bold>Kenya</bold>, from <italic>Delphinium</italic> sp.,
1990, Hopman, L 993.034.225, culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=177.93&link_type=cbs">CBS 177.93</ext-link> = PD
90/115. <bold>The Netherlands</bold>, Ter Aar, from <italic>Delphinium</italic> sp., 1986,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=176.93&link_type=cbs">CBS 176.93</ext-link> = PD
86/547.</p><p><italic><bold>Stagonosporopsis andigena</bold></italic> (Turkenst.) Aveskamp, Gruyter
& Verkley, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515655&link_type=mb">MB515655</ext-link>.</p><p><italic>Basionym</italic>: <italic>Phoma andigena</italic> Turkenst., apud Boerema, Gruyter
& Noordeloos, Persoonia 16(1): 131. 1995.</p><p><italic>Specimens examined</italic>: <bold>Peru</bold>, Dep. Junin, Huancayo, near Vallis
Mantaro, from a leaf of <italic>Solanum</italic> sp., 1975, L.J. Turkensteen,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=101.80&link_type=cbs">CBS 101.80</ext-link> = PD
75/909 = IMI 386090; Dep. Junin, Huancayo, near Vallis Mantaro, from a leaf of
<italic>Solanum</italic> sp., 1975, L.J. Turkensteen,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=269.80&link_type=cbs">CBS 269.80</ext-link> = PD
75/914.</p><p><italic><bold>Stagonosporopsis artemisiicola</bold></italic> (Hollós) Aveskamp,
Gruyter & Verkley, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515656&link_type=mb">MB515656</ext-link>.</p><p><italic>Basionym</italic>: <italic>Phoma artemisiicola</italic> Hollós, Mat.
Természettud. Közlem. 35: 40. 1926. (as
“<italic>artemisaecola</italic>”)</p><p><italic>Specimen examined</italic>: <bold>France</bold>, from a stem base of
<italic>Artemisia dracunculus</italic>, 1973,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=102636&link_type=cbs">CBS 102636</ext-link> = PD
73/1409.</p><p><italic><bold>Stagonosporopsis astragali</bold></italic> (Cooke & Harkn.) Aveskamp,
Gruyter & Verkley, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515657&link_type=mb">MB515657</ext-link>.</p><p><italic>Basionym</italic>: <italic>Phoma astragali</italic> Cooke & Harkn., Grevillea
13: 111. 1885.</p><p><italic>Specimen examined</italic>: Unknown origin, from <italic>Astragalus</italic> sp.,
1925, A.W. Archer, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=178.25&link_type=cbs">CBS
178.25</ext-link> = MUCL 9915.</p><p><italic><bold>Stagonosporopsis caricae</bold></italic> (Sydow & P. Sydow) Aveskamp,
Gruyter & Verkley, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515658&link_type=mb">MB515658</ext-link>.</p><p><italic>Basionym</italic>: <italic>Mycosphaerella caricae</italic> Sydow & P. Sydow,
Ann.. Mycol. 11: 403. 1913.</p><p><list list-type="simple"><list-item><p>≡ <italic>Phoma caricae-papayae</italic> (Tarr.) Punith., Trans Brit. Mycol.
Soc. 75: 340. 1980.</p></list-item><list-item><p>≡ <italic>Ascochyta caricae-papayae</italic> Tarr., The fungi and plant
diseases of Sudan: 53. 1955.</p></list-item><list-item><p>= <italic>Ascochyta caricae</italic> Pat., Bull. Soc. Mycol. France 7: 178.
1891.</p></list-item><list-item><p>≡ <italic>Phoma caricae</italic> Punith., CMI Descriptions of Pathogenic
Fungi and Bacteria 634: 1. 1979.</p></list-item></list></p><p>For description of the teleomorph see Sivanesan
(<xref ref-type="bibr" rid="ref207">1990</xref>). Punithalingam
(<xref ref-type="bibr" rid="ref170">1979b</xref>) provides an extensive
description, <italic>Ph. caricae</italic> is a synonym of the anamorph.</p><p><italic>Specimens examined</italic>: <bold>Brazil</bold>, from <italic>Carica papaya</italic>,
2006, J. de Gruyter, PD 06/03082531. <bold>Chile</bold>, from fruit of <italic>Carica
papaya</italic>, Feb. 1990, H.A. van der Aa,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=248.90&link_type=cbs">CBS 248.90</ext-link>.
<bold>Indonesia</bold>, Java, Segunung, from <italic>Brassica</italic> sp., Feb. 1976, H.
Vermeulen, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=282.76&link_type=cbs">CBS
282.76</ext-link>.</p><p><italic>Notes</italic>: <italic>Phoma caricae-papayae</italic> has been associated with an
undescribed teleomorph state in <italic>Mycosphaerella</italic> or <italic>Didymella</italic>(<xref ref-type="bibr" rid="ref55">Boerema <italic>et al.</italic>2004</xref>). Sivanesan
(<xref ref-type="bibr" rid="ref207">1990</xref>) synonymised <italic>Ph.
caricae</italic> with <italic>M. caricae</italic>, apparently not noting that <italic>Ph.
caricae</italic> already was synonymised with <italic>Ph. caricae-papayae</italic> by
Punithalingam (<xref ref-type="bibr" rid="ref171">1980</xref>). As
<italic>Mycosphaerella</italic> is phylogenetically unrelated to <italic>Phoma</italic>(<xref ref-type="bibr" rid="ref100">De Gruyter <italic>et al.</italic>2009</xref>), this taxonomic association is unlikely, and the observed
sexual state observed was probably <italic>Didymella</italic>-like.</p><p>This species has solely been associated with pawpaw (<italic>Carica papaya,
Caricaceae</italic>), but a single strain, deposited at CBS as <italic>D. exigua</italic>and that was isolated from <italic>Brassica</italic> leaves from Java, Indonesia
(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=282.76&link_type=cbs">CBS 282.76</ext-link>), was
genetically identical to the reference strain of <italic>Ph. caricae-papayae</italic>.
Herbarium material of this strain consisted of an inoculated lupine stem on
cornmeal agar (CBS H-11960) and represented a conidial state similar to this
of <italic>Ph. caricae-papayae</italic>. This indicated that probably the
<italic>Didymella</italic> teleomorph had been observed, but that it was preserved
under an incorrect name as it was only distantly related to the ex-type strain
of <italic>Didymella exigua</italic>(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=183.55&link_type=cbs">CBS 183.55</ext-link>). This
finding provides evidence that <italic>S. caricae</italic> is not restricted to
pawpaw.</p><p><italic><bold>Stagonosporopsis crystalliniformis</bold></italic> (Loer., R. Navarro, M.
Lôbo & Turkenst.) Aveskamp, Gruyter & Verkley, <bold>comb. nov.</bold>MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515659&link_type=mb">MB515659</ext-link>.</p><p><italic>Basionym</italic>: <italic>Phoma andina</italic> var. <italic>crystalliniformis</italic>Loer., R. Navarro, M. Lôbo & Turkenst., Fitopatología 21(2):
100. 1986.</p><p><list list-type="simple"><list-item><p>≡ <italic>Phoma crystalliniformis</italic> (Loer., R. Navarro, M. Lôbo
& Turkenst.) Noordel. & Gruyter, apud Noordeloos, de Gruyter, van Eijk
& Roeijmans, Mycol. Res. 97: 1344. 1993.</p></list-item></list></p><p><italic>Specimens examined</italic>: <bold>Colombia</bold>, Antioquia, Rionegro, from a
stem base of <italic>Lycopersicon esculentum</italic>, 1983, R. Navarro,
<bold>holotype</bold> CBS H-3926, ex-holotype culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=713.85&link_type=cbs">CBS 713.85</ext-link> = ATCC
76027 = PD 83/826; Guachacal, from a leaf of <italic>Solanum tuberosum</italic>, Nov.
1985, W.M. Loerakker, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=771.85&link_type=cbs">CBS
771.85</ext-link> = IMI 386091 = PD 85/772.</p><p><italic><bold>Stagonosporopsis cucurbitacearum</bold></italic> (Fr.) Aveskamp, Gruyter
& Verkley, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515660&link_type=mb">MB515660</ext-link>.</p><p><italic>Basionym</italic>: <italic>Sphaeria cucurbitacearum</italic> Fr., Syst. Mycol.
2(2): 502. 1823.</p><p><list list-type="simple"><list-item><p>≡ <italic>Phoma cucurbitacearum</italic> (Fr.) Sacc., Syll. Fung. 3: 148.
1884.</p></list-item><list-item><p>= <italic>Sphaeria bryoniae</italic> Fuck., Jahrb. Nassauischen Vereins Naturk.
23–24: 112. 1870.</p></list-item><list-item><p>≡ <italic>Didymella bryoniae</italic> (Fuckel) Rehm, Ber. Naturhist. Vereins
Augsburg 26: 27. 1881.</p></list-item></list></p><p><italic>Specimens examined</italic>: <bold>New Zealand</bold>, from <italic>Cucumis</italic> sp.,
1979, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=133.96&link_type=cbs">CBS 133.96</ext-link>=
PD 79/127. <bold>The Netherlands</bold>, Horst, from <italic>Cucumis</italic> sp., 1991, J.
de Gruyter, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=109171&link_type=cbs">CBS
109171</ext-link> = PD 91/310.</p><p><italic>Note</italic>: Strain <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=133.96&link_type=cbs">CBS
133.96</ext-link> could not be identified morphologically, as it proved to be
sterile.</p><p><italic><bold>Stagonosporopsis dennisii</bold></italic> Boerema, Gruyter & Noordel.,
Persoonia 16(3): 350. 1997.</p><p><list list-type="simple"><list-item><p>= <italic>Phoma dennisii</italic> Boerema, Trans. Brit. Mycol. Soc. 67(2): 307.
1976.</p></list-item></list></p><p><italic>Specimens examined</italic>: <bold>Canada</bold>, Ontario, from a stem of
<italic>Solidago canadensis</italic>, 1995, G.P. White,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=135.96&link_type=cbs">CBS 135.96</ext-link> = IMI
19337 = PD 94/4756. <bold>The Netherlands</bold>, Arnhem, from a stem of
<italic>Solidago floribunda</italic>, 1968,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=631.68&link_type=cbs">CBS 631.68</ext-link> = PD
68/147.</p><p><italic><bold>Stagonosporopsis dorenboschii</bold></italic> (Noordel. & Gruyter)
Aveskamp, Gruyter & Verkley, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515661&link_type=mb">MB515661</ext-link>.</p><p><italic>Basionym</italic>: <italic>Phoma dorenboschii</italic> Noordel. & Gruyter,
Persoonia 15(1): 83. 1992.</p><p><italic>Specimens examined</italic>: <bold>The Netherlands</bold>, Rijnsburg, from
<italic>Physostegia virginiana</italic>, 1986, D. Kruger, <bold>holotype</bold> L
988.202.121, <bold>isotype</bold> CBS H-7604, ex-holotype culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=426&link_type=cbs">CBS 426</ext-link>. 90 = IMI
386093 = PD 86/551; from <italic>Physostegia virginiana</italic>, 1986,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=320.90&link_type=cbs">CBS 320.90</ext-link> = PD
86/932.</p><p><italic><bold>Stagonosporopsis heliopsidis</bold></italic> (H.C. Greene) Aveskamp,
Gruyter & Verkley, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515662&link_type=mb">MB515662</ext-link>.</p><p><italic>Basionym</italic>: <italic>Phyllosticta heliopsidis</italic> H.C. Greene, Trans.
Wisconsin Acad. Sci. 50: 158. 1961.</p><p><list list-type="simple"><list-item><p>≡ <italic>Phoma heliopsidis</italic> (H.C. Greene), Aa & Boerema apud De
Gruyter Boerema & van der Aa, Persoonia 18(1): 40. 2002.</p></list-item></list></p><p><italic>Specimens examined</italic>: <bold>Canada</bold>, Island of Montréal, from
<italic>Ambrosia artemisiifolia</italic>, PD 95/6189 = DAOM 221138. <bold>The
Netherlands</bold>, from <italic>Heliopsis patula</italic>, 1974,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=109182&link_type=cbs">CBS 109182</ext-link> = PD
74/231.</p><p><italic><bold>Stagonosporopsis hortensis</bold></italic> (Sacc. & Malbr.) Petr., Ann.
Mycol. 19(1/2): 21. 1921.</p><p><italic>Basionym</italic>: <italic>Hendersonia hortensis</italic> Sacc. & Malbr., in
Saccardo, Michelia 2(8): 629. 1882.</p><p><list list-type="simple"><list-item><p>= <italic>Phoma subboltshauserii</italic> Boerema, Gruyter & Noordel.,
Persoonia 16(3): 360. 1997.</p></list-item></list></p><p><italic>Specimens examined</italic>: <bold>The Netherlands</bold>, from an unknown
substrate, Mar. 1942, N. Hubbeling,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=104.42&link_type=cbs">CBS 104.42</ext-link>; from
<italic>Phaseolus vulgaris</italic>, 1979, G.H. Boerema,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=572.85&link_type=cbs">CBS 572.85</ext-link> = PD
79/269.</p><p><italic><bold>Stagonosporopsis ligulicola</bold></italic> var.
<italic><bold>inoxydabilis</bold></italic> (Boerema) Aveskamp, Gruyter & Verkley,
<bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515664&link_type=mb">MB515664</ext-link>.</p><p><italic>Basionym</italic>: <italic>Didymella ligulicola</italic> var. <italic>inoxydabilis</italic>Boerema, Stud. Mycol. 32: 10. 1990.</p><p><italic>Anamorph</italic>: <italic>Phoma ligulicola</italic> var. <italic>inoxydabilis</italic>Boerema, Stud. Mycol. 32: 10. 1990.</p><p><italic>Specimens examined</italic>: <bold>The Netherlands</bold>, from <italic>Chrysanthemum
parthenii</italic>, 1981, G.H. Boerema, <bold>holotype</bold> CBS H-7611, culture
ex-holotype <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=425.90&link_type=cbs">CBS
425.90</ext-link> = PD 81/520; from <italic>Matricaria</italic> sp. 1985, J. de
Gruyter, PD 85/259.</p><p><italic><bold>Stagonosporopsis ligulicola</bold></italic> var. <italic><bold>ligulicola</bold></italic>(K.F. Baker, Dimock & L.H. Davis) Aveskamp, Gruyter & Verkley,
<bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515663&link_type=mb">MB515663</ext-link>.</p><p><italic>Basionym</italic>: <italic>Mycosphaerella ligulicola</italic> K.F. Baker, Dimock
& L.H. Davis, Phytopathology 39: 799. 1949.</p><p><italic>Anamorph</italic>: <italic>Phoma ligulicola</italic> var. <italic>ligulicola</italic>Boerema, Stud. Mycol. 32: 9. 1990.</p><p><italic>Specimens examined</italic>: <bold>Germany</bold>, Berlin, from <italic>Chrysanthemum
indicum</italic>, 1963, R. Schneider, CBS H-11952, culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=500.63&link_type=cbs">CBS 500.63</ext-link> = MUCL
8090. <bold>The Netherlands</bold>, near Lisse, from a leaf of <italic>Chrysanthemum
indicum</italic>, 1984, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=137.96&link_type=cbs">CBS
137.96</ext-link> = PD 84/75.</p><p><italic><bold>Stagonosporopsis loticola</bold></italic> (Died.) Aveskamp, Gruyter &
Verkley, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515665&link_type=mb">MB515665</ext-link>.</p><p><italic>Basionym</italic>: <italic>Phoma loticola</italic> Died., Kryptog.-Fl. Mark
Brandenburg. 9, Pilze 7(1): 152. 1912.</p><p><italic>Specimens examined</italic>: <bold>New Zealand</bold>, Auckland, Mt. Albert, from
<italic>Lotus pedunculatus</italic>, 1981, P.R. Johnston, <bold>isotype</bold> CBS H-7612,
ex-isotype culture <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=562.81&link_type=cbs">CBS
562.81</ext-link> = PDDCC 6884; Auckland, from the stem of <italic>Lotus
tenuis</italic>, 1979, P.R. Johnston,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=628.97&link_type=cbs">CBS 628.97</ext-link> = PD
79/72.</p><p><italic><bold>Stagonosporopsis lupini</bold></italic> (Boerema & R. Schneid.)
Boerema, Gruyter & P. Graaf, Persoonia 17(2): 283. 1999.</p><p><italic>Basionym</italic>: <italic>Ascochyta lupini</italic> Boerema & R. Schneid.,
apud Boerema, Verslagen Meded. Plziektenk. Dienst Wageningen 162: 28.
1984.</p><p><list list-type="simple"><list-item><p>≡ <italic>Phoma schneiderae</italic> (Boerema & R. Schneid.) Boerema,
Gruyter & P. Graaf, Persoonia 17(2): 282. 1999.</p></list-item></list></p><p><italic>Specimens examined</italic>: <bold>Peru</bold>, Puno, from <italic>Lupinus
mutabilis</italic>, 1980, CBS H-9061, culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=375.84&link_type=cbs">CBS 375.84</ext-link>= PD
80/1250. <bold>U.K.</bold>, Cambridgeshire, Mepal, from <italic>Lupinus albus</italic>, Apr.
1998, P. van de Graaf, <bold>holotype</bold> L 998.099.105, ex-holotype culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=101494&link_type=cbs">CBS 101494</ext-link> = PD
98/5247.</p><p><italic><bold>Stagonosporopsis oculo-hominis</bold></italic> (Punith.) Aveskamp, Gruyter
& Verkley, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515666&link_type=mb">MB515666</ext-link>.</p><p><italic>Basionym</italic>: <italic>Phoma oculi-hominis</italic> Punith., Trans Brit. Mycol.
Soc. 67: 142. 1976.</p><p><list list-type="simple"><list-item><p>≡ <italic>Phoma dennisii</italic> var. <italic>oculo-hominis</italic> (Punith.)
Boerema, Gruyter & Noordel., Persoonia 16: 351. 1997.</p></list-item></list></p><p><italic>Specimen examined</italic>: <bold>U.S.A.</bold>, Tennessee, Nashville, from a
man's corneal ulcer, 23 Apr. 1975, Y.M. Clayton, ex-holotype culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=634.92&link_type=cbs">CBS 634.92</ext-link> = IMI
193307.</p><p>For a complete description see Punithalingam
(<xref ref-type="bibr" rid="ref168">1976</xref>) and Boerema <italic>et
al.</italic> (<xref ref-type="bibr" rid="ref54">1997</xref>).</p><p><italic>Notes</italic>: <italic>Stagonosporopsis oculo-hominis</italic> is a species that
thus far has been reported only once in a clinical case in Tennessee, U.S.A.,
when it was isolated from a man's corneal ulcer
(<xref ref-type="bibr" rid="ref168">Punithalingam 1976</xref>). Due to
morphological similarities it has been recombined into a variety of <italic>Ph.
dennisii</italic> by Boerema <italic>et al.</italic>(<xref ref-type="bibr" rid="ref54">1997</xref>), but the genetical data
presented here suggest that this entity should be recognised at species level
in <italic>Stagonosporopsis</italic>. It is distinguishable from <italic>S. dennisii</italic>by the absence of a diffusible pigment in the agar, and by the absence of a
discolouration after application of NaOH to the culture. Further, the septate
conidia are significantly smaller than those of <italic>S. dennisii</italic>:
9–16 × 4.5 μm versus 14.5–24 × 4–7 μm,
respectively.</p><p><italic><bold>Stagonosporopsis rudbeckiae</bold></italic> (Fairm.) Aveskamp, Gruyter
& Verkley, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515667&link_type=mb">MB515667</ext-link>.</p><p><italic>Basionym</italic>: <italic>Phoma rudbeckiae</italic> Fairm., Proc. Rochester Acad.
Sci. 1: 51. 1890.</p><p><italic>Specimen examined</italic>: <bold>The Netherlands</bold>, from <italic>Rudbeckia
bicolor</italic>, 1979, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=109180&link_type=cbs">CBS
109180</ext-link> = PD 79/175.</p><p><italic><bold>Stagonosporopsis trachelii</bold></italic> (Allesch.) Aveskamp, Gruyter
& Verkley, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515668&link_type=mb">MB515668</ext-link>.</p><p><italic>Basionym</italic>: <italic>Phoma trachelii</italic> Allesch., Fungi Bavaria exs. 4:
360. 1897.</p><p><list list-type="simple"><list-item><p>= <italic>Ascochyta bohemica</italic> Kabát & Bubák apud
Bubák & Kabát, Hedwigia 44: 361. 1905.</p></list-item><list-item><p>≡ <italic>Stagonosporopsis bohemica</italic> (Kabát &
Bubák) Boerema, Gruyter & Noordel., Persoonia 16(3): 361. 1997.</p></list-item></list></p><p><italic>Specimens examined</italic>: <bold>Sweden</bold>, Svalöv, from <italic>Campanula
isophylla</italic>, 1968, W. Södergren, CBS H-8972, ex-holotype culture
384.68. <bold>The Netherlands</bold>, from a leaf of <italic>Campanula isophylla</italic>,
1977, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=379.91&link_type=cbs">CBS 379.91</ext-link> =
PD 77/675.</p><p><italic>Notes</italic>: Although this species has been described in
<italic>Stagonosporopsis</italic> before (as <italic>S. bohemica</italic>,
<xref ref-type="bibr" rid="ref54">Boerema <italic>et al.</italic> 1997</xref>),
this was based on a later homonym, and thus a recombination based on the
oldest epithet is proposed here.</p><p><italic><bold>Stagonosporopsis valerianellae</bold></italic> (Gindrat, Semecnik &
Bolay) Aveskamp, Gruyter & Verkley, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515669&link_type=mb">MB515669</ext-link>.</p><p><italic>Basionym</italic>: <italic>Phoma valerianellae</italic> Gindrat, Semecnik &
Bolay, Revue Hort. Suisse Romande 40: 350. 1967.</p><p><italic>Specimens examined</italic>: <bold>The Netherlands</bold>, Wageningen, from
<italic>Valerianella locusta</italic> var. <italic>oleracea</italic>, 1966, G.H. Boerema,
<bold>holotype</bold> L 965.300.24, <bold>isotype</bold> CBS H-7631, ex-isotype culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=329.67&link_type=cbs">CBS 329.67</ext-link> = PD
66/302; from <italic>Valerianella locusta</italic>, 1982,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=273.92&link_type=cbs">CBS 273.92</ext-link> = PD
76/1019.</p></sec><sec><title>Residual species in the Didymellaceae:</title><p>The following <italic>Phoma</italic> species are embedded in the
<italic>Didymellaceae</italic>, but could not be confidently assigned to one of the
groups or new genera in this study due to lack of support for their respective
clades. Several of the species listed here belong to this family based on LSU
and/or ITS sequence data, but due to missing sequencing data on one of the
loci used, these species could not be assigned. These species are
provisionally retained under their current holomorph name until further
analyses are conducted to place them in the new phylogenetic system.</p><p><italic><bold>Didymella macropodii</bold></italic> Petr., <italic>Hedwigia</italic> <bold>68</bold>:
219. 1928.</p><p><italic>Anamorph</italic>: <italic>Phoma nigrificans</italic> (P. Karst.) Boerema, Loer.
& Wittern, J. Phytopathol. 115(3): 270. 1986.</p><p><italic>Basionym</italic>: <italic>Sphaeronaema nigrificans</italic> P. Karst., Meddeland.
Soc. Fauna Fl. Fenn. 16: 17. 1888. (as
“<italic>Sphaeronema</italic>”).</p><p><italic>Specimens examined</italic>: <bold>Germany</bold>, from <italic>Brassica napus</italic>,
1982, G.H. Boerema, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=100190&link_type=cbs">CBS
100190</ext-link> = PD 82/736. <bold>Poland</bold>, near Gryfice, from <italic>Thlaspi
arvense</italic>, 1990, J. Marcinkowska,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=100191&link_type=cbs">CBS 100191</ext-link>. <bold>The
Netherlands</bold>, from an unidentified crucifer, 1984, G.H. Boerema, PD
84/512.</p><p><italic><bold>Didymella rabiei</bold></italic> (Kovatsch.) Arx, in Müller & Arx,
Beitr. Kryptogamenfl. Schweiz 11(2): 364. (1962).</p><p><italic>Basionym</italic>: <italic>Mycosphaerella rabiei</italic> Kovatsch., The blight of
chick pea: 70. 1936.</p><p><italic>Anamorph</italic>: <italic>Phoma rabiei</italic> (Pass.) Khune ex Gruyter,
Persoonia 18(1): 89. 2002.</p><p><italic>Basionym</italic>: <italic>Zythia rabiei</italic> Pass., Comment. Soc. Crittog.
Ital. 2(3): 437. 1867.</p><p><italic>Specimens examined</italic>: <bold>India</bold>, from the seeds of <italic>Cicer
arietinum</italic>, 1965, S. Sinha,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=534.65&link_type=cbs">CBS 534.65</ext-link>.
<bold>Syria</bold>, from <italic>Cicer arietinum</italic>, 1981, W. Barz,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=581.83&link_type=cbs">CBS 581.83</ext-link>A.</p><p><fig position="float" id="fig13"><label>Fig. 13.</label><caption><p><italic>Phoma bulgarica</italic>(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=357.84&link_type=cbs">CBS 357.84</ext-link>).
A–C. Fourteen-day-old colonies on OA (A), MEA (B) and CHA (C). D.
Pycnidia <italic>in vivo</italic>, isolated from manually infected sterilised stems of
<italic>Urtica dioica</italic>. E. Pycnidial section. F. Pycnidium. G. Crystals. H.
Hyphal strand. Scale bars: D, F = 100 μm; E, H = 50 μm; G = 10
μm.</p></caption><graphic xlink:href="1fig13"></graphic></fig></p><p><italic>Notes</italic>: The placement of this teleomorph in either the
<italic>Didymella</italic> or <italic>Mycosphaerella</italic> has been debated regularly in
the past (<xref ref-type="bibr" rid="ref143">Müller and Arx
1962</xref>, <xref ref-type="bibr" rid="ref15">Von Arx
1987</xref>, <xref ref-type="bibr" rid="ref224">Trapero-Casas &
Kaiser 1992</xref>, <xref ref-type="bibr" rid="ref236">Wilson &
Kaiser 1995</xref>, <xref ref-type="bibr" rid="ref94">De Gruyter
2002</xref>, <xref ref-type="bibr" rid="ref25">Barve <italic>et al.</italic>2003</xref>). The most recent emendment was by De Gruyter
(<xref ref-type="bibr" rid="ref94">2002</xref>) who judged in favour of
<italic>Mycosphaerella rabiei</italic> Kovatsch. ex Gruyter. However, as the genus
<italic>Mycosphaerella</italic> is phylogenetically not linked with the
<italic>Pleosporales</italic> (Schoch <italic>et al.</italic><xref ref-type="bibr" rid="ref193">2006</xref>,
<xref ref-type="bibr" rid="ref195">2009b</xref>,
<xref ref-type="bibr" rid="ref76">Crous <italic>et al</italic>. 2009c</xref>),
the placement in <italic>Didymella</italic> appears to be more correct.</p><p><italic><bold>Didymella adianticola</bold></italic> Aa & Boerema, Verslagen Meded.
Plziektenk. Dienst Wageningen 159 (Jaarboek 1982): 25. 1983.</p><p><italic>Anamorph</italic>: <italic>Phoma adianticola</italic> (E. Young) Boerema,
<italic>V</italic>erslagen Meded. Plziektenk. Dienst Wageningen 159 (Jaarboek 1982):
25. 1983</p><p><italic>Basionym</italic>: <italic>Phyllosticta adianticola</italic> E. Young, Mycologia 7:
144. 1915.</p><p><italic>Specimens examined</italic>: <bold>Costa Rica</bold>, from a leaf of
<italic>Polystichum adiantiforme</italic>, 1989, J. de Gruyter,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=258.92&link_type=cbs">CBS 258.92</ext-link> = PD
89/1887. <bold>U.S.A.</bold>, Florida, from a leaf of <italic>Polystichum
adiantiforme</italic>, 1982, G.H. Boerema, CBS H-16142, culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=187.83&link_type=cbs">CBS 187.83</ext-link> = PD
82/128.</p><p><italic><bold>Phoma aliena</bold></italic> (Fr.) Aa & Boerema, apud Gruyter,
Noordeloos & Boerema, Persoonia 16(4): 486. 1998.</p><p><italic>Basionym</italic>: <italic>Sphaeria aliena</italic> Fr., Syst. Mycol. 2(2): 502.
1823.</p><p><italic>Specimens examined</italic>: <bold>The Netherlands</bold>, from a twig of
<italic>Berberis</italic> sp., 1982, J. de Gruyter,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=379.93&link_type=cbs">CBS 379.93</ext-link> = PD
82/945; near Boskoop, from a twig of <italic>Buxus sempervirens</italic>, 1994, J. de
Gruyter, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=877.97&link_type=cbs">CBS 877.97</ext-link>= PD 94/1401.</p><p><italic><bold>Phoma aubrietiae</bold></italic> (Moesz) Boerema, Gewasbescherming 1(4):
66. 1970.</p><p><italic>Basionym</italic>: <italic>Sclerophomella aubrietiae</italic> Moesz,
Balkán-Kutat Tud. Eredm. 3: 144. 1926.</p><p><italic>Specimens examined</italic>: <bold>The Netherlands</bold>, Bodegraven, from seed
of <italic>Aubrietia hybrida</italic> cv. Superbissima, 1965, G.H. Boerema, CBS
H-16154, culture <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=383.67&link_type=cbs">CBS
383.67</ext-link> = PD 65/223; from a stem of <italic>Aubrietia</italic> sp., 1970,
G.H. Boerema, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=627.97&link_type=cbs">CBS
627.97</ext-link> = PD 70/714.</p><p><italic><bold>Phoma bulgarica</bold></italic> Aveskamp, Gruyter & Verkley, <bold>sp.
nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515671&link_type=mb">MB515671</ext-link>.
<xref ref-type="fig" rid="fig13">Fig. 13</xref>.</p><p>Pycnidia solitaria, subglobosa, elongata vel obpyriformia, glabra,
epapillata, brunnea, superficialia vel in agaro immersa,
(140–)170–250(–295) μm. Pycnidia fertilia non vidi.</p><p><italic>Etymology</italic>: Epithet refers to the country of origin, Bulgaria.</p><p><italic>Conidiomata</italic> pycnidial solitary, subglobose to elongated or
obpyriform, glabrous, non-papillate, brown, on the surface and immersed in the
agar, measuring (140–)170–250(–295) μm. Pycnidia proved
to be sterile. In older cultures pycnidial primordia are formed, which are
surrounded by clusters of needle-shaped crystals.</p><p><italic>Culture characteristics</italic>: Colonies on OA, 45–65 mm diam after
7 d, margin regular. Immersed mycelium hyaline, largely covered by mat of
felty to compact whitish grey to lavender grey aerial mycelium; reverse
iron-grey, but vinaceous-black where the aerial mycelium is present. Colonies
on MEA 40–50 mm diam after 7 d, margin regular. Immersed mycelium mainly
hyaline, incidentally black when clustering into thicker hyphal strands.
Aerial mycelium sparse, flat, olivaceous green to white near the colonies
margin; reverse greenish olivaceous to olivaceous black. Colonies on CHA
70–85 mm diam after 7 d, or even covering the total agar surface, margin
regular. Immersed mycelium as on MEA. Aerial mycelium occurring around the
colony centre, white, compact to floccose; reverse leaden black. Application
of NaOH did not have any effect.</p><p><italic>Specimens examined</italic>: <bold>Bulgaria</bold>, Silkossia, Strandga Mountain,
from leafs of <italic>Trachystemon orientale</italic>, 20 June 1980, S. Vanev,
<bold>holotype designated here</bold> CBS H-20242, ex-holotype culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=357.84&link_type=cbs">CBS 357.84</ext-link>; from
<italic>Trachystemon orientale</italic>, 1982,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=124515&link_type=cbs">CBS 124515</ext-link> = PD
82/1058.</p><p><italic>Notes</italic>: Strain PD 82/1058 differed from
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=357.84&link_type=cbs">CBS 357.84</ext-link> (which
is described above) by a significantly different colony pattern on MEA. This
strain was characterised by a growth of <italic>ca</italic>. 20 mm diam. after 7 d,
with a strongly lobate margin. White to buff aerial mycelium was present in a
few irregular zones, and had a compact to floccose structure. Pycnidial
primordial are only produced in culture on MEA after addition of an autoclaved
piece of <italic>Urtica dioica</italic> (stinging nettle).</p><p><italic><bold>Phoma calidophila</bold></italic> Aveskamp, Gruyter & Verkley,
Mycologia 101: 368. 2009.</p><p><italic>Specimens examined</italic>: <bold>Egypt</bold>, from desert soil, Feb. 1980,
M.I.A. Abdel-Kader, <bold>neotype</bold> CBS H-20168, ex-neotype culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=448.83&link_type=cbs">CBS 448.83</ext-link>. Unknown
European origin, from <italic>Cucumis sativus</italic>, 1984, G.H. Boerema, PD
84/109.</p><p><italic><bold>Phoma chenopodiicola</bold></italic> Gruyter, Noordel. & Boerema,
Persoonia 15(3): 395. 1993.</p><p><italic>Specimens examined</italic>: <bold>Peru</bold>, from a stem of <italic>Chenopodium
quinoa</italic> cv. Sajana, 1979,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=128.93&link_type=cbs">CBS 128.93</ext-link>= PD
79/140; from a stem of <italic>Chenopodium quinoa</italic> cv. Sajana, 1979,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=129.93&link_type=cbs">CBS 129.93</ext-link> = PD
89/803.</p><p><italic><bold>Phoma complanata</bold></italic> (Tode) Desm., <italic>Michelia</italic> <bold>2</bold>(7): 337. 1881. <italic>Basionym</italic>: <italic>Sphaeria complanata</italic> Tode, Fungi
Mecklenburg. Sel. (Lüneburg) 2: 22. 1791.</p><p><italic>Specimens examined</italic>: <bold>The Netherlands</bold>, Tilburg, from a stem
of <italic>Heracleum sphondylium</italic>, Nov. 1997, H.A. van der Aa, CBS H-16194,
culture <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=100311&link_type=cbs">CBS 100311</ext-link>;
from a stem of <italic>Angelica sylvestris</italic>, 1974, G.H. Boerema,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=268.92&link_type=cbs">CBS 268.92</ext-link> = PD
75/3.</p><p><italic><bold>Phoma crystallifera</bold></italic> Gruyter, Noordel. & Boerema,
Persoonia 15(3): 393. 1993.</p><p><italic>Specimen examined</italic>: <bold>Austria</bold>, Kärnten, Wallenberg near
Völkermarkt, from <italic>Chamaespartium sagittale</italic>, Apr. 1982, H.A. van
der Aa, <bold>holotype</bold> L 992.177-456, ex-holotype culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=193.82&link_type=cbs">CBS 193.82</ext-link>.</p><p><italic><bold>Phoma dactylidis</bold></italic> Aveskamp, Gruyter & Verkley, <bold>sp.
nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515671&link_type=mb">MB515671</ext-link>.
<xref ref-type="fig" rid="fig14">Fig. 14</xref>.</p><p>Conidia dimorpha, intra idem pycnidium formata. Conidia typus 1 ellipsoidea
vel ovoidea, interdum leniter allantoidea, glabra, hyalina, continua,
4.5–9(–9) × (2–)2.5–3.5 μm,
(2–)3–6(–8) guttulis praedita. Conidia typus 2 cylindrica
vel ellipsoidea, glabra, hyalina, saepe uniseptata,
(9–)9.5–13.5(–14.5) × (2.5–)3.5–4.5 μm,
interdum septata et guttulis (2–)4–8(–15) in quoque cellula.
Matrix conidiorum salmonea.</p><p><italic>Etymology</italic>: Named after the associated plant host genus,
<italic>Dactylis</italic> sp.</p><p><italic>Conidiomata</italic> pycnidial, solitary or confluous, produced on the agar
surface, (sub-)globose, with some hyphal outgrows,
(115–)135–230(–250) ×
(75–)95–195(–105) μm. <italic>Ostioles</italic>1–4(–5), papillate. <italic>Pycnidial wall</italic> pseudoparenchymatous,
composed of isodiametric cells, 4–8 cell layers, outer 2–4 cell
layers pigmented, 10–27 μm thick. <italic>Conidiogenous cells</italic>phialidic, hyaline, simple, smooth, flask-shaped, <italic>ca.</italic> 4.5–6.5
× 3–5 μm. <italic>Conidia</italic> dimorphic, both originating from the
same pycnidia. Conidia of type 1: ellipsoidal to ovoid, sometimes somewhat
allantoid, thin-walled, smooth, hyaline, aseptate 4.5–9(–9)
× (2–)2.5–3.5 μm, with (2–)3–6(–8)
guttules. Regularly also large conidia occur: cylindrical to ellipsoidal,
thin-walled, smooth, hyaline, often uniseptate
(9–)9.5–13.5(–14.5) × (2.5–)3.5–4.5 μm,
but sometimes septate and septate somewhat constricted at the septum, with
(2–)4–8(–15) guttules per cell. Conidial matrix salmon.</p><p><italic>Culture characteristics</italic>: Colonies on OA, 40–45 mm diam after
7 d, margin regular. Immersed mycelium hyaline, but some greenish black zones
may occur, with tufts of with aerial mycelium. Abundant greenish black
pycnidia are scattered over the medium, which are salmon coloured near the
colony margin; reverse concolourous. Colonies on MEA 45–50 mm diam after
7 d, margin regular. Immersed mycelium completely covered by a felty greyish
white aerial mycelium; reverse grey-olivaceous, becoming brown-olivaceous near
the colony margin. Colonies on CHA similar as on MEA, but somewhat faster
growing, 55–60 mm diam. after 7 d; reverse completely black. Application
of NaOH results in a slight greenish discolouration of the agar, best to be
observed on OA medium.</p><p><italic>Specimen examined</italic>: <bold>U.S.A.</bold>, Oregon, on <italic>Dactylis
glomerata</italic>, 1973, <bold>holotype designated here</bold> CBS H-20237, ex-holotype
culture <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=124513&link_type=cbs">CBS 124513</ext-link>= PD 73/1414.</p><p><italic>Notes</italic>: <italic>Phoma dactylidis</italic> has thus far only been isolated
once from the leaves of <italic>Dactylis glomerata</italic> in Oregon, U.S.A.. Other
<italic>Phoma</italic> pathogens of <italic>Dactylis</italic> include <italic>Ph. paspali</italic> and
<italic>Ph. pratorum,</italic> which both occur in New Zealand, but are relatively
distantly related to <italic>Ph. dactylidis</italic>. Additionally, two related taxa
have been found on this host, viz. the novel variety <italic>Boeremia exigua</italic>var. <italic>gilvescens</italic> and <italic>Epicoccum nigrum</italic>(<xref ref-type="bibr" rid="ref172">Punithalingam <italic>et al.</italic>1972</xref>). The clustering of this species suggests ecological or
morphological similarities with <italic>Ph. rhei</italic> (BPP = 1.00; RBS = 100
%).</p><p><italic><bold>Phoma destructiva</bold></italic> var. <italic><bold>destructiva</bold></italic> Plowr.,
Gard. Chron. II 16: 621. 1881.</p><p><italic>Specimens examined</italic>: <bold>Guadeloupe</bold>, from fruit of
<italic>Lycopersicon esculentum</italic>, 1987,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=133.93&link_type=cbs">CBS 133.93</ext-link> = PD
88/961 = IMI 173142. <bold>Tonga</bold>, Friendly Islands, from decaying fruit of
<italic>Lycopersicon esculentum</italic>, 1967, G.F. Laundon, CBS H-16200, culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=378.73&link_type=cbs">CBS 378.73</ext-link> = CECT
2877.</p><p><italic><bold>Phoma destructiva</bold></italic> var. <italic><bold>diversispora</bold></italic> Gruyter
& Boerema, apud De Gruyter, Boerema & Van der Aa, Persoonia 18(1): 28.
2002.</p><p><italic>Specimen examined</italic>: <bold>The Netherlands</bold>, Berkel en Rodenrijs,
from a leaf of <italic>Lycopersicon esculentum</italic>, Oct. 1977, G.H. Boerema,
<bold>holotype</bold> CBS H-16199, ex-holotype culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=162.78&link_type=cbs">CBS 162.78</ext-link> = PD
77/725.</p><p><italic><bold>Phoma eupyrena</bold></italic> Sacc., Michelia 1(5): 525. 1879.</p><p><italic>Specimens examined</italic>: <bold>Germany</bold>, Kiel-Kitzeberg, from wheat
field soil, 1966, W. Gams, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=527.66&link_type=cbs">CBS
527.66</ext-link> = ATCC 22238; <bold>The Netherlands</bold>, from the tuber of
<italic>Solanum tuberosum</italic>, 1991, J. de Gruyter,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=374.91&link_type=cbs">CBS 374.91</ext-link> = PD
78/391.</p><p><fig position="float" id="fig14"><label>Fig. 14.</label><caption><p><italic>Phoma dactylidis</italic>(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=124513&link_type=cbs">CBS 124513</ext-link>).
A–C. Fourteen-day-old colonies on OA (A), MEA (B) and CHA (C). D-E.
Pycnidia. F. Section of the pycnidial wall. G. Conidia. Scale bars: D–E
= 100 μm; F–G = 5 μm.</p></caption><graphic xlink:href="1fig14"></graphic></fig></p><p><italic><bold>Phoma herbarum</bold></italic> Westend., Bull. Acad. Roy. Sci. Belgique,
Cl. Sci. 19(3): 118. 1852.</p><p><italic>Specimens examined</italic>: <bold>Belgium</bold>, Herb. Crypt. Belge. Fasc. 20,
No. 965, <bold>lectotype</bold>, on stems of <italic>Onobrychis viciifolia</italic>, 1854.
Sweden, Sofieheim, from wood pulp, Apr. 1937, E. Rennerfelt,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=276.37&link_type=cbs">CBS 276.37</ext-link> = MUCL
9920. <bold>The Netherlands</bold>, Emmeloord, from the stem of <italic>Rosa
multiflora</italic> cv. Cathayensis, Apr. 1965, G.H. Boerema,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=615.75&link_type=cbs">CBS 615.75</ext-link> = PD
73/665 = IMI 199779; Naaldwijk, from a stem base of <italic>Nerium</italic> sp., 1986,
J. de Gruyter, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=502.91&link_type=cbs">CBS
502.91</ext-link> = PD 82/276; Oirschot, from a twig of <italic>Thuja</italic> sp.,
1987, J. de Gruyter, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=503&link_type=cbs">CBS
503</ext-link>. 91 = PD 87/499. <bold>U.K.</bold>, from paint, Aug. 1936, K.S.G.
Cartwright, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=109.36&link_type=cbs">CBS
109.36</ext-link>. <bold>U.S.A.</bold>, Maryland, Washington area, from the fruit
of <italic>Malus sylvestris</italic>, July 1963, M.A. Smith,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=567.63&link_type=cbs">CBS 567.63</ext-link> = ATCC
15053 = MUCL 9889.</p><p><italic><bold>Phoma herbicola</bold></italic> Wehm., Mycologia 38: 319. 1946.</p><p><italic>Specimen examined</italic>: <bold>U.S.A.</bold>, Montana, Missoula, head of
Seeley Lake, from water, CBS H-16581, culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=629.97&link_type=cbs">CBS 629.97</ext-link> = PD
76/1017.</p><p><italic><bold>Phoma huancayensis</bold></italic> Turkenst., Fitopatologia 13: 68.
1978.</p><p><italic>Specimens examined</italic>: <bold>Peru</bold>, Dep. Junin, Huancayo, near Vallis
Mantaro, from a stem of <italic>Solanum</italic> sp., Feb. 1974, L.J. Turkensteen,
<bold>isotype</bold> CBS H-7609, ex-isotype culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=105.80&link_type=cbs">CBS 105.80</ext-link> = PD
75/908; from <italic>Chenopodium quinoa</italic>, 1977,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=390.93&link_type=cbs">CBS 390.93</ext-link> = PD
77/1173.</p><p><italic><bold>Phoma longicolla</bold></italic> Aveskamp, Gruyter & Verkley, <bold>sp.
nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515672&link_type=mb">MB515672</ext-link>.
<xref ref-type="fig" rid="fig15">Fig. 15</xref>.</p><p>Conidia late ellipsoidea vel ovoidea, glabra, hyalina, continua,
6–8.5(–10) × (3.5–)4–5(–5.5) μm,
(2–)3–9(–12) guttulis polaris praedita. Matrix conidiorum
cremeo-alba.</p><p><italic>Etymology</italic>: Refers to the elongated necks of the ostioles.</p><p><italic>Conidiomata</italic> pycnidial, initially solitary, globose, glabrous,
slightly papillate and olivaceous buff, produced on the agar surface,
measuring (45–)50–115(–130) μm diam. Later developing to
black broadly globose to irregular conidiomata with many white hyphal outgrows
and with a clear elongated neck around the ostioles, giving it a irregular
shape, measuring (170–)200–270(–285) ×
(115–)125–205(–220) μm. <italic>Ostioles</italic>1–3(–4), on a long elongated neck (up to 200 μm long). Often
these pycnidia merge to an irregular mass of confluent conidiomata.
<italic>Pycnidial wall</italic> pseudoparenchymatous, composed of isodiametric cells,
5–7 layers, 17–22 μm thick. <italic>Conidiogenous cells</italic>phialidic, hyaline, simple, smooth, flask-shaped, <italic>ca</italic>. 4–5
× 3–5 μm. <italic>Conidia</italic> broadly ellipsoidal to ovoid,
thin-walled, smooth, hyaline, aseptate 6–8.5(–10) ×
(3.5–)4–5(–5.5) μm, with (2–)3–9(–12)
polar guttules. Conidial matrix crème-white.</p><p><fig position="float" id="fig15"><label>Fig. 15.</label><caption><p><italic>Phoma longicolla</italic>(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=124514&link_type=cbs">CBS 124514</ext-link>).
A–C. Fourteen-day-old colonies on OA (A), MEA (B) and CHA (C).
D–F. Pycnidia. G. Section of the pycnidial wall. H. Conidia. Scale bars:
E–G = 100 μm; H = 50 μm; I–J = 10 μm.</p></caption><graphic xlink:href="1fig15"></graphic></fig></p><p><italic>Culture characteristics</italic>: Colonies on OA 50–55 mm diam. after
7 d, margin regular. Immersed mycelium hyaline with abundant pycnidia, in some
sectors covered by a low mat of felty to floccose mouse grey aerial mycelium,
with tufts of white mycelium near the colonies margin. In the sectors with
aerial mycelium, pycnidia are only sparsely present; reverse hyaline, but
leaden black and olivaceous grey where the aerial mycelium is present. Colony
on MEA 50–55 mm diam. after 7 d, margin regular. Immersed mycelium
completely covered by a floccose crème mat of white aerial mycelium;
reverse greenish olivaceous to olivaceous-black. Colony on CHA 55–60 mm
diam. after 7 d, margin regular. Immersed mycelium brown vinaceous to black.
Aerial mycelium is occurring in sectors, felty, pale grey to white; reverse
black with incidentally a pale purplish grey zone. Application of NaOH did not
have any effect.</p><p><italic>Specimens examined</italic>: <bold>Spain</bold>, Canary Isles, from
<italic>Opuntia</italic> sp., 1980, J. de Gruyter, <bold>holotype designated here</bold> CBS
H-20238, ex-holotype culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=124514&link_type=cbs">CBS 124514</ext-link> = PD
80/1189; Canary Isles, Gran Canaria, from <italic>Opuntia</italic> sp., June 1982,
H.A. van der Aa, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=347.82&link_type=cbs">CBS
347.82</ext-link>.</p><p><italic>Notes</italic>: This species was isolated twice from <italic>Opuntia</italic> on
the Canary Isles. Around the time of the second isolation
(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=347.82&link_type=cbs">CBS 347.82</ext-link>), also
<italic>Ph. dimorpha</italic> sp. nov. was isolated from the same location and host
substrate. This species is described above. A third species that is found in
association with <italic>Opuntia</italic> is <italic>Ph. opuntiae</italic>, which is, however,
rather distinct in morphology and phylogeny.</p><p><italic><bold>Phoma medicaginis</bold></italic> var. <italic><bold>macrospora</bold></italic> Boerema,
R. Pieters & Hamers, Netherlands J. Pl. Pathol. 99(Suppl. 1): 19.
1993.</p><p><italic>Specimens examined</italic>: <bold>Canada</bold>, Saskatchewan, Saskatoon, from
seed of <italic>Medicago sativa</italic>, 1965, H.W. Mead,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=404.65&link_type=cbs">CBS 404.65</ext-link> = IMI
116999. <bold>U.S.A.</bold>, Minnesota, from <italic>Medicago sativa</italic>, Sep. 1953,
M.F. Kernkamp, <bold>holotype</bold> CBS H-16487, ex-holotype culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=112.53&link_type=cbs">CBS 112.53</ext-link>.</p><p><italic><bold>Phoma medicaginis</bold></italic> var. <italic><bold>medicaginis</bold></italic> Malbr.
& Roum. apud Roumeguère, Fungi Selecti Galliaei Exs. 37: 3675.
1886.</p><p><italic>Specimens examined</italic>: <bold>Czech Republic</bold>, from <italic>Medicago
sativa</italic>, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=316.90&link_type=cbs">CBS
316.90</ext-link> = CCM F-187. <bold>Italy</bold>, Perugia, from a leaf of
<italic>Medicago sativa</italic>, 1963, M. Ribaldi, CBS H-16483, culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=479.63&link_type=cbs">CBS 479.63</ext-link>. <bold>The
Netherlands</bold>, from a leaf of <italic>Medicago sativa</italic>, 1966, M.M.J.
Dorenbosch, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=533.66&link_type=cbs">CBS
533.66</ext-link> = ATCC 16929 = PD 66/370. <bold>Turkey</bold>, Ankara, from
<italic>Medicago sativa</italic>, 1942, S. Kuntay,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=107.42&link_type=cbs">CBS 107.42</ext-link>.
<bold>U.S.A.</bold>, Minnesota, from <italic>Medicago sativa</italic>, Sep. 1953, M.F.
Kernkamp, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=110.53&link_type=cbs">CBS
110.53</ext-link>; Minnesota, from <italic>Medicago sativa</italic>, Sep. 1953, M.F.
Kernkamp, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=111.53&link_type=cbs">CBS
111.53</ext-link>.</p><p><italic><bold>Phoma microchlamydospora</bold></italic> Aveskamp & Verkley, Mycologia
101: 374. 2009.</p><p><italic>Specimens examined</italic>: <bold>U.K.</bold>, from an unknown vegetable plant,
1990, D. Hyall, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=491.90&link_type=cbs">CBS
491.90</ext-link>; from leaves of <italic>Eucalyptus</italic> sp., 1994, A.M.
Ainsworth, <bold>holotype</bold> CBS H-20147, ex-holotype culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=105.95&link_type=cbs">CBS 105.95</ext-link>.</p><p><italic><bold>Phoma nebulosa</bold></italic> (Pers.) Berk., <italic>Outl. Brit. Fung.
(London)</italic>: 314. 1860.</p><p><italic>Basionym</italic>: <italic>Sphaeria nebulosa</italic> Pers., Observ. Mycol. 2: 69.
1799.</p><p><italic>Specimens examined</italic>: <bold>Austria</bold>, Kaprun, from a stem of
<italic>Urtica dioica</italic>, Jan. 1975, G.H. Boerema, CBS H-16510, culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=503.75&link_type=cbs">CBS 503.75</ext-link>= ATCC
32163 = DSM 63391 = IMI 194766 = PD 75/4. <bold>The Netherlands</bold>, from a stem
of <italic>Mercurialis perennis</italic>, 1983,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=117.93&link_type=cbs">CBS 117.93</ext-link> = PD
83/90.</p><p><italic><bold>Phoma negriana</bold></italic> Thüm., Die Pilze des Weinstockes,
Vienna: 185. 1878. Originally described as “<italic>Ph.
negrianum</italic>”.</p><p><italic>Specimens examined</italic>: <bold>Germany</bold>, Oberdollendorf am Rhein, from
<italic>Vitis vinifera</italic>, July 1969, L. Kiewnik, CBS H-16511, culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=358.71&link_type=cbs">CBS 358.71</ext-link>. <bold>The
Netherlands</bold>, from <italic>Vitis vinifera</italic>, 1979, PD 79/74; from <italic>Vitis
vinifera</italic>, 1979, PD 79/75; from <italic>Vitis vinifera</italic>, 1979, PD
79/76.</p><p><italic><bold>Phoma nigripycnidia</bold></italic> Boerema, Gruyter & Noordel.,
Persoonia 16(3): 356. 1997.</p><p><italic>Specimen examined</italic>: <bold>Russia</bold>, from a leaf of <italic>Vicia
cracca</italic>, 1969, M. Ondrej, <bold>holotype</bold> L 992.163.150, ex-holotype
culture <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=116.96&link_type=cbs">CBS 116.96</ext-link>= CCMF 243 = PD 95/7930.</p><p><italic><bold>Phoma omnivirens</bold></italic> Aveskamp, Verkley & Gruyter, Mycologia
101: 375. 2009.</p><p><italic>Specimens examined</italic>: <bold>Belgium</bold>, Gembloux, from <italic>Phaseolus
vulgaris</italic>, 1968, L. Obando, <bold>holotype</bold> CBS H-20151, ex-holotype
culture <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=341.86&link_type=cbs">CBS 341.86</ext-link>.
<bold>India</bold>, Japalbur, from an unknown substrate, 1977, D.P. Tiwari,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=654.77&link_type=cbs">CBS 654.77</ext-link>.
<bold>Papua New Guinea</bold>, Varirata National Park, from soil, Aug. 1995, A.
Aptroot, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=991.95&link_type=cbs">CBS
991.95</ext-link>. Varirata National Park. From soil, Aug. 1995, A. Aptroot,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=992.95&link_type=cbs">CBS 992.95</ext-link>.
<bold>Tanzania</bold>, from <italic>Statice</italic> sp., 1990, J. de Gruyter,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123397&link_type=cbs">CBS 123397</ext-link> = PD
90/1555. <bold>The Netherlands</bold>, from <italic>Chrysanthemum indicum</italic>, 1981, J.
de Gruyter, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123396&link_type=cbs">CBS
123396</ext-link> = PD 81/122.</p><p><italic><bold>Phoma putaminum</bold></italic> Speg., Atti Soc. Crittog. Ital. 3: 66.
1881.</p><p><italic>Specimens examined</italic>: <bold>The Netherlands</bold>, from a branch of
<italic>Ulmus</italic> sp., 1975, G.H. Boerema,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=372.91&link_type=cbs">CBS 372.91</ext-link> = PD
75/960. <bold>Denmark</bold>, from the rhizosphere of <italic>Malus sylvestris</italic>,
Mar. 1968, E. Sønderhousen,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=130.69&link_type=cbs">CBS 130.69</ext-link> = CECT
20054 = IMI 331916.</p><p><italic><bold>Phoma rhei</bold></italic> (Ellis & Everh.) Aa & Boerema apud De
Gruyter, Boerema & Van der Aa, Persoonia 18 (1): 42. 2002.</p><p><italic>Basionym</italic>: <italic>Ascochyta rhei</italic> Ellis & Everh., Proc. Acad.
Nat. Sci. Philadelphia 1893: 160. 1893.</p><p><italic>Specimen examined</italic>: <bold>New Zealand</bold>, from a leaf of <italic>Rheum
rhaponticum</italic>, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=109177&link_type=cbs">CBS
109177</ext-link> = LEV 15165 = PD 2000/9941.</p><p><italic><bold>Phoma selaginellicola</bold></italic> Gruyter, Noordel., Aa & Boerema,
Persoonia 15(3): 399. 1993.</p><p><italic>Specimen examined</italic>: <bold>The Netherlands</bold>, from a leaf of
<italic>Selaginella</italic> sp., 1977, G.H. Boerema,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=122.93&link_type=cbs">CBS 122.93</ext-link> = PD
77/1049.</p><p><italic><bold>Phoma versabilis</bold></italic> Boerema, Loer. & Hamers, Persoonia
16(2): 154. 1996.</p><p><italic>Specimens examined</italic>: <bold>Germany</bold>, Westfalen, Oberdresselendorf,
from stems of <italic>Cardamine impatiens</italic>, Oct. 1925, A. Ludwig,
<bold>holotype</bold> L 995.229.369. <bold>The Netherlands</bold>, Wageningen, from a stem
of <italic>Silene</italic> sp., 1982, G.H. Boerema,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=876&link_type=cbs">CBS 876</ext-link>. 97 = PD
82/1008; from <italic>Stellaria media</italic>, 2000, J. de Gruyter, PD 2000/1379.</p></sec></sec></sec></sec><sec><title>DISCUSSION</title><sec><title>What is <italic>Phoma</italic>?</title><p>According to the generic concept which is applied today, species of
<italic>Phoma</italic> are relatively simple coelomycetes that are characterised by
the <italic>in vitro</italic> production of mainly unicellular, hyaline conidia from
monophialidic, doliiform to flask-shaped conidiogenous cells in pycnidial
conidiomata (<xref ref-type="bibr" rid="ref36">Boerema & Bollen
1975</xref>).</p><p>Many species that currently are accommodated in sections <italic>Paraphoma,
Pilosa</italic> and <italic>Plenodomus</italic> are phylogenetically basal to the
<italic>Didymellaceae</italic>, in which most other <italic>Phoma</italic> taxa, including the
type species are accommodated. These results support the work of Reddy <italic>et
al.</italic> (<xref ref-type="bibr" rid="ref178">1998</xref>), who
advocated that the genus <italic>Plenodomus</italic> should be reinstalled as a
separate genus. Torres <italic>et al.</italic>(<xref ref-type="bibr" rid="ref226">2005b</xref>) subsequently made a
novel description in this genus, <italic>Pl. morganjonesii</italic>. A paper by De
Gruyter <italic>et al.</italic> is in preparation, in which all species of
<italic>Phoma</italic> section <italic>Plenodomus</italic> recognised by Boerema <italic>et
al.</italic> (<xref ref-type="bibr" rid="ref52">1994</xref>,
<xref ref-type="bibr" rid="ref53">1996</xref>) and Boerema & De
Gruyter (<xref ref-type="bibr" rid="ref41">1999</xref>), will be
taxonomically revised.</p><p>However, in the present study, it has become clear that the phylogenetic
boundaries between <italic>Phoma</italic> and several closely related genera that are
defined on their conidial characters are ambiguous. Species that produce
consistently two-celled hyaline conidia classified therefore traditionally in
the genus <italic>Ascochyta</italic> appear to have evolved independently multiple
times during evolution together with typical <italic>Phoma</italic> taxa, in several
lineages of the pleosporalean tree (<xref ref-type="fig" rid="fig1">Fig.
1</xref>). Also other conidial characters, such as the pigmentation of
spores, as formed by <italic>Phoma clematidis-rectae</italic> (formerly in
<italic>Coniothyrium</italic>) and <italic>Microsphaeropsis olivacea,</italic> appear not to
be reliable for the delimitation of the genus <italic>Phoma</italic>. Thus, based on
the trees presented in this study, it can be concluded that <italic>Phoma</italic>, as
defined by Saccardo (<xref ref-type="bibr" rid="ref187">1880</xref>,
<xref ref-type="bibr" rid="ref188">1884</xref>) and emended by Boerema
& Bollen (<xref ref-type="bibr" rid="ref36">1975</xref>) is highly
polyphyletic.</p><p>The close relation of <italic>Phoma</italic> with <italic>Ascochyta</italic> has often been
observed before, as strains of both genera are often highly similar in
morphology (Wollenweber & Hochapfel 1937,
<xref ref-type="bibr" rid="ref57">Brewer & Boerema 1965</xref>,
<xref ref-type="bibr" rid="ref36">Boerema & Bollen 1975</xref>,
<xref ref-type="bibr" rid="ref34">Boerema 1997</xref>), physiology
(<xref ref-type="bibr" rid="ref152">Noordeloos <italic>et al.</italic>1993</xref>, Faris-Mokaiesh <italic>et al.</italic> 1995), pathogenicity
(Mendes-Perreira <italic>et al.</italic> 1999,
<xref ref-type="bibr" rid="ref80">Davidson <italic>et al.</italic> 2009</xref>)
and nucleotide sequences (Faris-Mokaiesh <italic>et al.</italic> 1995,
<xref ref-type="bibr" rid="ref88">Fatehi <italic>et al.</italic> 2003</xref>,
<xref ref-type="bibr" rid="ref193">Schoch <italic>et al.</italic> 2006</xref>,
<xref ref-type="bibr" rid="ref163">Peever <italic>et al.</italic> 2007</xref>,
<xref ref-type="bibr" rid="ref66">Chilvers <italic>et al.</italic> 2009</xref>,
<xref ref-type="bibr" rid="ref100">De Gruyter <italic>et al.</italic>2009</xref>). In the Saccardoan system, both genera were only
distinguished by the presence or absence of conidial septa, and by the type of
substrate: <italic>Ascochyta</italic> species were considered to be specific
leaf-pathogens, whereas <italic>Phoma</italic> was solely associated with stem lesions
(<xref ref-type="bibr" rid="ref36">Boerema & Bollen
1975</xref>).</p><p>Brewer & Boerema (<xref ref-type="bibr" rid="ref57">1965</xref>)
contrasted the septation process of the conidia in <italic>Ascochyta pisi</italic> to
this process in <italic>Phoma exigua</italic>. These authors suggested that in
<italic>Phoma</italic> species euseptation occurs only after secession, whereas in
<italic>Ascochyta</italic> the septation of the spores was considered to be an
elemental part of conidiogenesis. Later, this was determined to be a
genus-specific character (<xref ref-type="bibr" rid="ref31">Boerema
1970</xref>). Additionally, Boerema and Bollen
(<xref ref-type="bibr" rid="ref36">1975</xref>) stated that both genera
are distinct in conidiogenesis. According to these authors, the
<italic>Ascochyta</italic> species produce conidia from either an accumulation of
annelations, which give the conidigenous cell an annelidic appearance, or from
a gradually thickening collar of periclinal annelations. In contrast,
<italic>Phoma</italic> species produce true phialides with a collarette. This
micromorphological difference of the conidiogenesis can only be observed using
electron microscopy, as the appearance of a <italic>Phoma</italic> collarette is
highly similar to the periclinal thickening of <italic>Ascochyta</italic> species.
This observation is however not consistent with the conidial ontogeny of
<italic>Ph. fumosa</italic>, which was observed to be annellidic by Sutton &
Sandhu (<xref ref-type="bibr" rid="ref220">1969</xref>).</p><p>The application of these characters for the purpose of generic delimitation
was heavily questioned (<xref ref-type="bibr" rid="ref169">Punithalingam
1979a</xref>), and nowadays these characters are hardly applied in the
taxonomy of both genera, simply because the use of electron microscopy is
expensive and sectioning of pycnidia is too time consuming. Due to this
unclear classification system, and to the fact that not all species produce
exclusively septate or aseptate conidia, species had synonyms in both genera
(<xref ref-type="bibr" rid="ref32">Boerema 1972</xref>,
<xref ref-type="bibr" rid="ref37">Boerema & Dorenbosch 1973</xref>,
<xref ref-type="bibr" rid="ref7">Van der Aa <italic>et al.</italic>2000</xref>, <xref ref-type="bibr" rid="ref134">Mel'nik
2000</xref>). Even nowadays the status of many species is unclear as
<italic>Phoma</italic> and <italic>Ascochyta</italic> synonyms are often used simultaneously.
Examples are <italic>Ph. rabiei</italic> and its synonym <italic>A. rabiei</italic>(<xref ref-type="bibr" rid="ref205">Singh & Reddy 1993</xref>,
<xref ref-type="bibr" rid="ref206">Singh <italic>et al.</italic> 1997</xref>,
<xref ref-type="bibr" rid="ref25">Barve <italic>et al.</italic> 2003</xref>,
<xref ref-type="bibr" rid="ref67">Chongo <italic>et al.</italic> 2004</xref>,
<xref ref-type="bibr" rid="ref158">Pande <italic>et al.</italic> 2005</xref>,
<xref ref-type="bibr" rid="ref106">Hernandez-Bello <italic>et al.</italic>2006</xref>, <xref ref-type="bibr" rid="ref163">Peever <italic>et al.</italic>2007</xref>), and <italic>Ph. gossypiicola</italic> and its synonym, <italic>A.
gossypii</italic> (<italic>e.g.</italic> <xref ref-type="bibr" rid="ref200">Shen <italic>et
al.</italic> 2005</xref>). The concept of <italic>Ph. clematidina</italic> has
appeared to comprise several taxa belonging in multiple genera, amongst which
a <italic>Didymella</italic> with an unnamed <italic>Ascochyta</italic> anamorph
(<xref ref-type="bibr" rid="ref239">Woudenberg <italic>et al.</italic>2009</xref>).</p><p>The results presented in this study further suggest a close relation
between <italic>Microsphaeropsis</italic> and <italic>Phoma</italic>. Morphological studies of
members of both genera (<xref ref-type="bibr" rid="ref121">Jones
1976</xref>) reveal that conidiogenesis is similar, although the conidia
of <italic>Microsphaeropsis</italic> differ from those of <italic>Phoma</italic> by the dark
pigmentation and the presence of a double-layered cell wall. The pigmentation
occurs only after conidial secession. Therefore, young pycnidia with
colourless pycnidia may be easily confused with a <italic>Phoma</italic> species
(<xref ref-type="bibr" rid="ref55">Boerema <italic>et al</italic>.
2004</xref>).</p><p>In general, it can be concluded that <italic>Phoma</italic> should only be regarded
as a general concept, as members sharing this morphology are found throughout
the <italic>Pleosporales</italic>, although most members are found in the
<italic>Didymellaceae</italic>. The type species of <italic>Phoma</italic> is only distantly
related to the other members of this genus, but relatively close to
<italic>Ascochyta pisi</italic>, the type species of the older name
<italic>Ascochyta</italic>. However, based on the results observed in the present
study, this genus is poorly elucidated. Therefore, we opt to retain the
taxonomy of <italic>Phoma</italic> as is, with the exception of the groups that can be
resolved further, such as <italic>Boeremia, Epicoccum, Peyronellaea</italic> and
<italic>Stagonosporopsis</italic>.</p></sec><sec><title>Taxonomic revisions</title><p>The observations presented in the present paper suggest that LSU and SSU
data, which contain approximately 270 informative sites in the alignment, are
sufficient to distinguish various major groups in the <italic>Pleosporales</italic>.
However, other, more variable loci should also be analysed to determine the
phylogenetical basis for the species that are congeneric with the ex-type
strain of <italic>Phoma</italic>. These species were found throughout the
pleosporalean phylogeny that was reconstructed in the present paper. Molecular
studies on the species that are currently accommodated in the section
<italic>Plenodomus</italic> and <italic>Pilosa</italic> are in progress (De Gruyter <italic>et
al.</italic> in prep.).</p><p>The type species of the genus <italic>Phoma, Ph. herbarum,</italic> resides in the
<italic>Didymellaceae</italic> clade, a result that is in congruence with the
observations of De Gruyter <italic>et al.</italic>(<xref ref-type="bibr" rid="ref100">2009</xref>). However, based on the
data generated in the present study, also the type species of
<italic>Ascochyta</italic> (<italic>A. pisi</italic>), <italic>Chaetasbolisia</italic> (<italic>C.
erysiphoides</italic>), and <italic>Microsphaeropis</italic> (<italic>M. olivaceae</italic>) are
located in the same group (<xref ref-type="fig" rid="fig2">Fig.
2</xref>). Of those species, <italic>Phoma</italic> carries the oldest name,
which was deposited by Fries in 1821, but as <italic>Phoma sensu</italic> Saccardo
(<xref ref-type="bibr" rid="ref187">1880</xref>) was conserved
against<italic> Phoma</italic> Fries (<xref ref-type="bibr" rid="ref133">McNeill
<italic>et al.</italic> 2006</xref>), the genus <italic>Ascochyta</italic>, which was
erected in 1830, would be the preferred name for the species in this genus.
Nevertheless, because of the impact that recombination of <italic>Phoma</italic> in
<italic>Ascochyta</italic> would have in phytopathology, we suggest to keep both
generic names in use for the unresolved species in the <italic>Didymellaceae</italic>,
disregarding the fact that both names are polyphyletic. Both genera can be
regarded as polyphyletic concepts, until a proper study of the teleomorph
genera related to the <italic>Didymellaceae</italic> has been conducted. Also the
younger genera <italic>Chaetasbolisia</italic>and <italic>Microsphaeropsis</italic> should be
retained as separate taxonomic entities, until at least all taxa are restudied
both morphologically and phylogenetically. However, the clades that are
resolved, and that are characterised by shared morphological or physiological
characters, or have a shared ecological role, are elevated to generic level
here. Consequences of this approach are the reinstatement of the genus
<italic>Peyronellaea</italic> Goid., expansion of the formerly monotypic genus
<italic>Epicoccum</italic> Link, emendment of the concept of <italic>Stagonosporopsis</italic>Died. and the erection of the novel genus <italic>Boeremia</italic>.</p></sec><sec><title>Teleomorph relations</title><p>In <italic>Phoma</italic> several teleomorphs have been recognised, but for the
majority of <italic>Phoma</italic> species the sexual structures have yet to be
discovered, as the induction of these structures requires special conditions;
or simply because the species has lost its ability to propagate sexually.
Boerema <italic>et al.</italic> (<xref ref-type="bibr" rid="ref55">2004</xref>)
only recognised <italic>ca</italic>. 40 species that produce teleomorphs.</p><p>The finding of multiple teleomorphs with phenotypically indistinguishable
associated anamorphs is not uncommon in mycology, yet unwanted, and should be
resolved in due course as more data become available. For example, such a
situation also applies to major genera such as <italic>Aspergillus</italic>(<xref ref-type="bibr" rid="ref166">Pitt & Samson 2007</xref>),
<italic>Botryosphaeria</italic> (<xref ref-type="bibr" rid="ref73">Crous <italic>et
al.</italic> 2006</xref>), <italic>Geotrichum</italic>(<xref ref-type="bibr" rid="ref109">De Hoog & Smith 2004</xref>),
<italic>Mycosphaerella</italic> (Crous <italic>et al.</italic><xref ref-type="bibr" rid="ref74">2009a</xref>,
<xref ref-type="bibr" rid="ref75">b</xref>) and <italic>Penicillium</italic>(<xref ref-type="bibr" rid="ref165">Pitt 1979</xref>).</p><p>Boerema <italic>et al.</italic>(<xref ref-type="bibr" rid="ref55">2004</xref>) linked <italic>Phoma</italic>to four teleomorph genera: <italic>Didymella, Leptosphaeria, Mycosphaerella</italic>and <italic>Pleospora</italic>. In recent studies it was shown that the association of
<italic>Phoma</italic> with <italic>Mycosphaerella</italic> was untenable, because the
involved teleomorphs were apparently morphologically similar but in fact
<italic>Didymella</italic>. The genus <italic>Mycosphaerella</italic> is phylogenetically
distinct and not even associated with the <italic>Pleosporales</italic> (Schoch <italic>et
al.</italic> <xref ref-type="bibr" rid="ref193">2006</xref>,
<xref ref-type="bibr" rid="ref194">2009a</xref>,
<xref ref-type="bibr" rid="ref195">b</xref>, Crous <italic>et al</italic>.
<xref ref-type="bibr" rid="ref74">2009a</xref>,
<xref ref-type="bibr" rid="ref75">b</xref>), whereas their associated
<italic>Phoma</italic> anamorphs proved to be genetically similar to
<italic>Didymella</italic> (<xref ref-type="bibr" rid="ref100">De Gruyter <italic>et
al.</italic> 2009</xref>). As a consequence, the pawpaw (<italic>Carica
papaya</italic>) pathogen <italic>M. caricae</italic> has been recombined into <italic>D.
caricae</italic> in the present study. Also the <italic>Didymellaceae</italic> clade is
not yet completely resolved.</p><p>Next to <italic>Didymella</italic>, also <italic>Leptosphaerulina</italic> and
<italic>Macroventuria</italic> are accommodated in the <italic>Didymellaceae.
Macroventuria</italic> resembles <italic>Venturia</italic> (Van der Aa <italic>et al.</italic>1971); the ascospore morphology being highly comparable to that of
<italic>Didymella</italic>. In contrast, <italic>Leptosphaerulina</italic> is distinct in
morphology, producing ascospores with longitudinal and transverse septa, more
resembling the ascospores of <italic>Pleospora</italic> and <italic>Cucurbitaria</italic>(<xref ref-type="bibr" rid="ref14">Von Arx 1981</xref>).
<italic>Didymella</italic> is a poorly studied genus that is in need of a
comprehensive revision, as it plays such a crucial role in the delimitation of
phytopathologically important genera. When studied more intensively, this
genus may very well be split up into multiple genera that have a proper
morphological basis.</p><p>Sexual states have thus far only been reported for a limited number of
<italic>Phoma</italic> species. It seems unlikely that the ability to produce sexual
reproductive structures is lost in so many species, whilst other, closely
related species, or even species that emerge from these “asexual”
species, do have a teleomorph state. It may be assumed that the sexual state
of these species is cryptic, and can only be induced under the right
conditions. These teleomorph structures, that probably much resemble the
sexual structures formed by the genus <italic>Didymella</italic>, are probably the
missing links that are required for further taxonomical delineation of the
species in the <italic>Didymellaceae</italic>.</p></sec><sec><title>Can the sections be maintained?</title><p>The present study was initiated chiefly to clarify the status of
<italic>Phoma</italic> and to judge the validity of the sections introduced by Boerema
(<xref ref-type="bibr" rid="ref34">1997</xref>). Aveskamp <italic>et
al.</italic> (<xref ref-type="bibr" rid="ref18">2008</xref>) already
illustrated the ambiguity of some sections, as multiple characters that are
regarded to be section-specific may be present in a single species. For
example, <italic>Ph. zeae-maydis</italic> was regarded as the type species of the
section <italic>Macrospora</italic>, due to the presence of its relatively large
aseptate spores (<xref ref-type="bibr" rid="ref94">De Gruyter
2002</xref>). However, this species also produces multicellular
chlamydospores, resembling the chlamydospores formed in species that are
accommodated in the section <italic>Peyronellaea</italic>. The recombination of this
species into <italic>Pey. zeae-maydis</italic> in the present study, which is based on
DNA phylogeny, indicates that the spore size is not an informative character
at above-species level.</p><p>Another example of the ambiguity of the Boeremaean section is <italic>Ph.
destructiva</italic>. Infraspecific taxa of this species are accommodated in two
sections: <italic>Ph. destructiva</italic> var. <italic>diversispora</italic> was accommodated
in section <italic>Phyllostictoides</italic>, wheras the type variety was linked to
section <italic>Phoma</italic> due to the absence of septate conidia. Boerema <italic>et
al.</italic> (<xref ref-type="bibr" rid="ref55">2004</xref>) acknowledged
this ambiguity problem and were forced to key out several species in multiple
sectional dichotomous keys. In the previous study of De Gruyter <italic>et
al</italic>. (<xref ref-type="bibr" rid="ref100">2009</xref>) this
ambiguity could not be illustrated as only sectional representatives were
included. Here it is illustrated that, although some sections can be partially
maintained, most of the sections are not supported from an evolutionary
perspective.</p><sec><title>Section <italic>Heterospora</italic></title><p>The majority of the species that were ascribed to <italic>Phoma</italic> section
<italic>Heterospora</italic> is recovered in Group R, from which the species are all
recombined into the genus <italic>Stagonosporopsis</italic> in the present paper. The
type species of section <italic>Heterospora</italic> however, <italic>Ph.
heteromorphospora</italic>, is recovered basal to the <italic>Didymellaceae</italic>together with <italic>Ph. dimorphospora</italic>. Also <italic>Ph. samarorum</italic> is not
retrieved in the main <italic>Phoma</italic> clade, but is associated with the
<italic>Phaeosphaeriaceae</italic>.</p><p>Also within the <italic>Didymellaceae</italic>, the <italic>Heterospora</italic> section
appears to be polyphyletic as <italic>Ph. aquilegiicola, Ph. glaucii</italic> and
<italic>Ph. clematidina</italic> are distantly related to most other
<italic>Heterospora</italic> species and form a distinct clade together with another
<italic>Clematidina</italic>pathogen, <italic>Ph. clematidis-rectae</italic>, a species that
has been regularly confused with the <italic>Phoma clematidina</italic> complex
(<xref ref-type="bibr" rid="ref239">Woudenberg <italic>et al.</italic>2009</xref>). The species in this clade can be distinguished from the
main body of the <italic>Heterospora</italic> species as they lack the production of
large <italic>Stagonospora</italic>-type conidia in culture, although smaller, septate
conidia may occur.</p></sec><sec><title>Section <italic>Macrospora</italic></title><p>The five large-spored species of the section <italic>Macrospora</italic> included
in this study are found scattered throughout the <italic>Didymellaceae</italic>,
indicating that spore size is not a good taxonomic criterion for delimiting
taxa above species level. <italic>Phoma zeae-maydis</italic> is genetically similar to
most <italic>Peyronellaea</italic> species. This association is supported by the
finding of dictyochlamydospores in most species in this clade
(<xref ref-type="bibr" rid="ref19">Aveskamp <italic>et al.</italic>2009a</xref>).</p></sec><sec><title>Section <italic>Paraphoma</italic></title><p>Also <italic>Phoma</italic> section <italic>Paraphoma</italic>(<xref ref-type="bibr" rid="ref6">Van der Aa <italic>et al.</italic>1990</xref>) appears to be polyphyletic. The section comprises 12 taxa
that produce pycnidial conidiomata with setae
(<xref ref-type="bibr" rid="ref95">De Gruyter & Boerema
2002</xref>). Members of this section are found in clades 5, 6, and 8 of
<xref ref-type="fig" rid="fig1">Fig. 1</xref>. <italic>Phoma gardenia</italic>is the only setae-producing species known in the <italic>Didymellaceae</italic>.
Because of its ability to produce dictyochlamydospores, and based on the DNA
phylogeny presented in <xref ref-type="fig" rid="fig2">Fig. 2</xref>,
it is recombined into the genus <italic>Peyronellaea</italic> here.</p><p>The type species for the former section <italic>Paraphoma</italic> is <italic>Ph.
radicina</italic>, which is accommodated in the <italic>Phaeosphaeriaceae</italic> group
(clade 6). Remarkably, no other species that were ascribed to the section
<italic>Paraphoma</italic> are found in the same family. Instead, <italic>Ph.
chrysanthemicola</italic> (formerly ascribed to the section <italic>Peyronellaea</italic>)
is found in close association with <italic>Ph. radicina</italic>. Both species are
recognised as soil fungi and have a wide distribution with records from
Europe, North-America and Asia (<xref ref-type="bibr" rid="ref55">Boerema
<italic>et al.</italic> 2004</xref>). The close association between <italic>Ph.
samarorum, Ph. chrysanthemicola</italic> and <italic>Ph. radicina</italic> has been
recorded before in a phylogenetical reconstruction of the section
<italic>Peyronellaea</italic> in a study of
<xref ref-type="bibr" rid="ref19">Aveskamp <italic>et al.</italic>2009a</xref>. The resolution of the clade in that study was, however,
higher as the complete ITS regions 1 and 2 were applied in genetic analyses
(<xref ref-type="bibr" rid="ref19">Aveskamp <italic>et al.</italic>2009a</xref>). Further linkage of the morphological and ecological
characters to the phylogeny will be one of the main challenges for taxonomists
working on the species in this group.</p><p>A third <italic>Paraphoma</italic> species, <italic>Ph. terricola</italic>, is recovered in
clade 5 of <xref ref-type="fig" rid="fig1">Fig. 1</xref>, which
resembles the <italic>Cucurbitaceae</italic>. This family also hosts the setae-lacking
species <italic>Ph. pratorum</italic>, which was classified in section <italic>Phoma</italic>.
Several other coelomycete fungi are accommodated here as well, including
<italic>Phialophorophoma litoralis, Pleurophoma cava</italic>, a sterile strain that
once has been identified as <italic>Coniothyrium</italic> sp. and various
<italic>Pyrenochaeta</italic> species. The close morphological relation between the
genera <italic>Pyrenochaeta, Pleurophoma</italic> and <italic>Phoma</italic> section
<italic>Paraphoma</italic> was already noted by Boerema <italic>et al.</italic>(<xref ref-type="bibr" rid="ref53">1996</xref>) and Grondona <italic>et
al.</italic> (<xref ref-type="bibr" rid="ref92">1997</xref>). Like
<italic>Phialophorophoma litoralis</italic> and <italic>Pleurophoma cava,
Pyrenochaeta</italic> is characterised by the formation of elongated, filiform,
multiseptate conidiophores, a character that is however not found in the
various <italic>Phoma</italic> species embedded in this clade
(<xref ref-type="bibr" rid="ref100">De Gruyter <italic>et al.</italic>2009</xref>). A further delineation of the species associated with the
genera <italic>Pyrenochaeta</italic> and <italic>Pleurophoma</italic> and the <italic>Phoma</italic>section <italic>Paraphoma</italic> will be provided in a follow-up paper by De Gruyter
<italic>et al.</italic> (<xref ref-type="bibr" rid="ref101">2010</xref>).</p></sec><sec><title>Section <italic>Peyronellaea</italic></title><p>The chlamydospore-producing species have been treated before by Aveskamp
<italic>et al.</italic> (<xref ref-type="bibr" rid="ref19">2009a</xref>), who
revealed that also <italic>Phoma</italic> section<italic> Peyronellaea</italic> is artificial
from an evolutionary point of view. Most species, including the type <italic>Ph.
glomerata</italic>, cluster in group K of <xref ref-type="fig" rid="fig2">Fig.
2</xref>, along with many other (uni- and multicellular) chlamydospore
producing species. To be in accordance with the phylogenetic results, this
cluster is elevated to generic level, which is named after the section
<italic>Peyronellaea.</italic> A second group of species belonging to this section is
recovered in clade L, which groups species that produce botryoid or epicoccoid
dictyochlamydospores, including <italic>Epicoccum nigrum</italic>. Two species,
<italic>Ph. pimprina</italic> and <italic>Ph. sorghina</italic> are recombined into
<italic>Epicoccum</italic> here. Species that produce pseudoscleroid chlamydospores,
such as <italic>Ph. violicicola</italic> and <italic>Ph. chrysanthemicola</italic> were found
to cluster outside the <italic>Didymellaceae</italic>.</p></sec><sec><title>Section <italic>Phoma</italic></title><p>Species ascribed to <italic>Phoma</italic> section <italic>Phoma</italic> are retrieved in
practically all clades of the trees produced in the present study. This
supports the general idea that this section has been used as a
“waste-bin” for phomoid taxa that could not be placed in other
sections or genera due to the lack or presence of typical sectional
characters.</p><p>The type species of this section, and also of the genus as a whole, is
<italic>Ph. herbarum</italic> (<xref ref-type="bibr" rid="ref29">Boerema
1964</xref>). The reference strains of this species are accommodated
amongst the basal polytomous species of the <italic>Didymellaceae</italic>. This
suggests that it has branched off from most other members of this family in an
early phase of the development of the <italic>Didymellaceae</italic> and probably
evolved further without recombining with other taxa.</p><p>Although the description of <italic>Ph. crinicola</italic> is highly similar to
that of other species in the <italic>B. exigua</italic> clade presented in
<xref ref-type="fig" rid="fig2">Fig. 2</xref>, it has never been
recognised as such due to the absence of septate conidia. Nevertheless, the
remaining characters do not contradict with the description given for <italic>Ph.
exigua</italic> (<xref ref-type="bibr" rid="ref7">Van der Aa <italic>et al.</italic>2000</xref>). The pycnidia of <italic>Ph. crinicola</italic> usually carry a
single ostiole, but pycnidia are regularly observed lacking an apparent
ostiole. This may correspond with the ostiolar openings of many species found
within the <italic>exigua</italic> clade, which are often lined or filled with
papillate, hyaline cells.</p><p>Similar findings are <italic>Ph. aurea</italic> and <italic>Ph. nigricans</italic> in clade
K, which is mainly filled with chlamydospore-forming species that were
previously associated with the section <italic>Peyronellaea</italic>. Both species
were originally described from New Zealand
(<xref ref-type="bibr" rid="ref120">Johnston & Boerema 1981</xref>,
<xref ref-type="bibr" rid="ref97">De Gruyter <italic>et al.</italic>1993</xref>), but may be commonly present on the whole Australasian
continent (De Gruyter <italic>et al.</italic><xref ref-type="bibr" rid="ref97">1993</xref>,
<xref ref-type="bibr" rid="ref98">1998</xref>). Two other species,
belonging to section <italic>Phoma</italic>, but found in this clade are <italic>Ph.
anserina</italic> and <italic>Ph. eucalyptica</italic>. Both species produce swollen cells
in older cultures (<xref ref-type="bibr" rid="ref96">De Gruyter &
Noordeloos 1992</xref>), which may be an initial phase of chlamydospore
formation.</p><p>Fifteen species are phylogenetically only distantly related to the
<italic>Didymellaceae</italic>, and should therefore be excluded from the genus. These
species include the current <italic>Ph. apiicola, Ph. capitulum, Ph. fallens, Ph.
fimeti, Ph. flavescens, Ph. flavigena, Ph. glaucispora, Ph. haematocycla, Ph.
lini, Ph. minutispora, Ph. multipora, Ph. opuntiae, Ph. pratorum, Ph.
valerianae</italic>, and <italic>Ph. vasinfecta</italic>. The problem in recombining these
species is, however, the absence of characters that could link these taxa to a
specific genus. No teleomorphs are known in this group.</p></sec><sec><title>Section <italic>Phyllostictoides</italic></title><p>All taxa belonging to <italic>Phoma</italic> section <italic>Phyllostictoides</italic> are
retrieved in the <italic>Didymellaceae</italic> clade of
<xref ref-type="fig" rid="fig1">Fig. 1</xref> (Clade 8). This is
remarkable as this large section has been regarded, just like section
<italic>Phoma</italic>, to be a repository for all species that could not be
accommodated elsewhere. Nevertheless, within the <italic>Didymellaceae</italic> this
section falls apart as species occur in many distinct clades.</p><p>The major body of the <italic>Phyllostictoides</italic> species is retrieved in
group N, in which all <italic>Ph. exigua</italic>-related species and varieties are
found (<xref ref-type="bibr" rid="ref20">Aveskamp <italic>et al.</italic>2009b</xref>), as well as <italic>Ph. crinicola</italic> and <italic>Ph.
hedericola</italic>, which were associated with <italic>Phoma</italic> section
<italic>Phoma</italic>. A second group in which many <italic>Phyllostictoides</italic> taxa
cluster is clade R. This clade comprises many species of the former section
<italic>Heterospora,</italic> and several species that were excluded from this section
and transferred to <italic>Phyllostictoides</italic> by Boerema <italic>et al.</italic>(<xref ref-type="bibr" rid="ref54">1997</xref>), such as <italic>Ph.
cucurbitacearum</italic> and <italic>Ph. ligulicola</italic>.</p></sec><sec><title>Section <italic>Pilosa</italic></title><p>Only one of both members of the section Pilosa was included in the present
study. The type of this section, <italic>Ph. betae</italic>, produces a teleomorph in
<italic>Pleospora</italic>, a genus that is typified by <italic>Pl. herbarum</italic>. Both
species are related and are found in the <italic>Pleosporaceae</italic> and
<italic>Leptosphaeriaceae</italic> clade, although the genetic distance between these
species is significant. This finding illustrates the difficulties that are
experienced when delineating the <italic>Pleosporaceae</italic>(<xref ref-type="bibr" rid="ref82">Dong <italic>et al.</italic>1998</xref>).</p></sec><sec><title>Section <italic>Plenodomus</italic></title><p>Thus far the only section created by Boerema that still may be monophyletic
is the section <italic>Plenodomus</italic>, of which all the members are found in the
<italic>Leptosphaeriaceae</italic>. However, some species associated with other
sections, such as <italic>Ph. apiicola, Ph. valerianae, Ph. vasinfecta</italic>(section <italic>Phoma)</italic> and <italic>Ph. violicola</italic> (section
<italic>Peyronellaea</italic>) are also linked to this clade and are found to be
closely related to the <italic>Plenodomus</italic> species. The section
<italic>Plenodomus</italic> is associated with a <italic>Leptosphaeria</italic> teleomorph,
but for the aberrant <italic>Phoma</italic> states found in this clade, no teleomorphs
are known. Boerema <italic>et al.</italic>(<xref ref-type="bibr" rid="ref55">2004</xref>) mentioned five
<italic>Leptosphaeria</italic> species that produce <italic>Phoma</italic> anamorphs, but that
do not fit within the <italic>Plenodomus</italic> concept. These species, including
<italic>L. sacchari, L. haematitis, L. libanotis, L. purpurea</italic> and <italic>L.
weimeri</italic> were however not to our disposal, and were therefore not studied.
Apparently the genus <italic>Leptosphaeria</italic> produces multiple anamorphs.</p><p>Most taxonomic studies on the <italic>Leptosphaeriaceae</italic> reveal a
monophyletic group, although in these studies, only a limited number of
species, belonging to either <italic>Leptosphaeria</italic> or <italic>Phoma</italic> section
<italic>Plenodomus,</italic> have been included
(<xref ref-type="bibr" rid="ref139">Morales <italic>et al.</italic>1995</xref>, <xref ref-type="bibr" rid="ref178">Reddy <italic>et al.</italic>1998</xref>, <xref ref-type="bibr" rid="ref226">Torres <italic>et al.</italic>2005b</xref>). Other studies indicate that this genus is paraphyletic
(<xref ref-type="bibr" rid="ref82">Dong <italic>et al.</italic> 1998</xref>,
<xref ref-type="bibr" rid="ref62">Câmara <italic>et al.</italic>2002</xref>). Due to the inclusion of only two <italic>Leptosphaeria</italic>species in the present study, it cannot be unambiguously stated whether this
section is mono- or paraphyletic.</p><p>Both species included, <italic>L. maculans</italic> and <italic>L. biglobosa</italic>, are
assumed to represent a heterogeneous assemblage of cryptic taxa
(<xref ref-type="bibr" rid="ref112">Howlett <italic>et al</italic>.
2001</xref>, <xref ref-type="bibr" rid="ref136">Mendes-Pereira <italic>et
al</italic>. 2003</xref>, <xref ref-type="bibr" rid="ref24">Barrins <italic>et
al</italic>. 2004</xref>, <xref ref-type="bibr" rid="ref231">Voigt <italic>et
al</italic>. 2005</xref>). Although many recombinations have been made in the
past, this has obscured a proper understanding of <italic>Phoma</italic> section
<italic>Plenodomus</italic> and <italic>Leptosphaeria</italic>(<xref ref-type="bibr" rid="ref53">Boerema <italic>et al.</italic>1996</xref>). Due to the complexity of this group, we will attempt to
resolve its phylogeny in a separate paper (De Gruyter <italic>et al.</italic>prep.).</p></sec><sec><title>Section <italic>Sclerophomella</italic></title><p>The thickened, sclerotisised pycnidial wall, and the formation of poroid
pycnidial openings instead of an ostiole, are the main characters of
<italic>Phoma</italic> section <italic>Sclerophomella</italic>. These characters appear not to
reflect the evolutionary history of the genus. Only in group O, a cluster of
species is retrieved that is known for their ostiole absence, although not in
all species the thickened pycnidial wall is observed. Most other species
belonging to section <italic>Sclerophomella</italic> appear to be unrelated as they
have emerged from non-<italic>Sclerophomella</italic> multiple times during evolution.
Therefore these species are found scattered throughout the phylogeny of the
Pleosporales. The type species of this section is <italic>Ph. complanata</italic>,
which is found in the basal polytomy of the <italic>Didymellaceae</italic>.</p><p>Many of the morphological characters that were used by Boerema <italic>et
al.</italic> (<xref ref-type="bibr" rid="ref54">1997</xref>) to create an
infrageneric subdivision of <italic>Phoma</italic>, appear not to be evolutionary
informative when compared to sequence data. The main characters that were
applied to distinguish sections, like the thickness of the pycnidial walls,
chlamydospore structure and presence of <italic>Stagonosporopsis</italic> synanamorphs
are only of limited value. Several characters, such as percentage of septated
spores may be genetically driven, but are certainly also highly influenced by
the growth media and culturing conditions
(<xref ref-type="bibr" rid="ref175">Rai 2000</xref>). This has led to
much confusion surrounding the taxonomic placement of many species in either
<italic>Ascochyta</italic> or <italic>Phoma</italic>, such as <italic>A. rabiei</italic>(<italic>e.g.</italic> <xref ref-type="bibr" rid="ref25">Barve <italic>et al.</italic>2003</xref>, <xref ref-type="bibr" rid="ref158">Pande <italic>et al.</italic>2005</xref>, <xref ref-type="bibr" rid="ref163">Peever <italic>et al.</italic>2007</xref>) vs. <italic>Ph. rabiei</italic> (<italic>e.g.</italic><xref ref-type="bibr" rid="ref205">Singh & Reddy 1993</xref>,
<xref ref-type="bibr" rid="ref206">Singh <italic>et al.</italic> 1997</xref>,
<xref ref-type="bibr" rid="ref94">De Gruyter 2002</xref>).</p><p>In short, the Boeremaean sectional subdivision is hardly of any
evolutionary relevance, suggesting that future classification of taxonomic
novelties into these sections should be avoided. Nevertheless, the
morphological identification system that was developed based on this
subdivision (<xref ref-type="bibr" rid="ref55">Boerema <italic>et al.</italic>2004</xref>) is still applicable, as this system can be still aid in
morphological species recognition.</p></sec></sec><sec><title>DNA Barcoding</title><p>A further aim of this study was the development of species-specific DNA
barcodes for species of <italic>Phoma</italic>. The preferred DNA barcode region for
<italic>Fungi</italic> is ITS (Druzhininia <italic>et al.</italic> 2005,
<xref ref-type="bibr" rid="ref215">Summerbell <italic>et al</italic>.
2005</xref>, Seifert
<xref ref-type="bibr" rid="ref196">2008</xref>,
<xref ref-type="bibr" rid="ref197">2009</xref>). Cytochrome Oxidase I
(COI) was for a long time considered to be a good candidate gene for barcoding
fungi (<xref ref-type="bibr" rid="ref198">Seifert <italic>et al.</italic>2007</xref>, <xref ref-type="bibr" rid="ref149">Nguyen & Seifert
2008</xref>), although some recent studies indicate the variation between
copies within a single strain (<xref ref-type="bibr" rid="ref90">Geiser
<italic>et al.</italic> 2007</xref>, <xref ref-type="bibr" rid="ref91">Gilmore
<italic>et al.</italic> 2009</xref>). Also Aveskamp <italic>et al.</italic>(<xref ref-type="bibr" rid="ref20">2009b</xref>) found that the COI
locus was not robust, and thus far, COI barcodes have only been applied in an
oligonucleotide array identification system for <italic>Penicillium</italic> spp.
(<xref ref-type="bibr" rid="ref65">Chen <italic>et al.</italic> 2009</xref>).
The value of ITS as primary barcode region is, however, not sufficient to
delineate all taxa. Especially amongst the species clustered in clade N, which
represents the species that are associated to the <italic>Ph. exigua</italic> species
complex, ITS is not sufficient to distinguish the various species. This
finding is in congruence with results obtained in previous studies, in which
the ITS region has been applied in an attempt to distinguish the species
within the <italic>Ph. exigua</italic> complex but without success
(<xref ref-type="bibr" rid="ref132">MacDonald <italic>et al.</italic>2000</xref>, <xref ref-type="bibr" rid="ref8">Abeln <italic>et al.</italic>2002</xref>, Cullen <italic>et al.</italic> 2006). Nevertheless, the other taxa
included in this study have been found on long-branched clades, which are
mainly due to the variation in TUB and ITS sequences. Another locus that is
considered to be helpful for developing DNA barcodes, and which can
distinguish many more taxa in the <italic>Ph. exigua</italic> complex is the Actin
gene (<xref ref-type="bibr" rid="ref20">Aveskamp <italic>et al.</italic>2009b</xref>), which is sequenced with a primer combination developed by
Carbone & Kohn (<xref ref-type="bibr" rid="ref63">1999</xref>).
This locus has, however, not been included in the present study, as
infraspecific genetic variation, even within the <italic>Didymellaceae</italic>, was
too high to align the obtained sequences. Also Calmodulin and Translation
Elongation Factor 1-α loci have been tested, but none of the primers
combinations used (<xref ref-type="bibr" rid="ref63">Carbone & Kohn
1999</xref>) could guarantee successful amplification of all strains.</p><p>Observations and results presented here represent only a preliminary step
towards resolving questions related to the taxonomy of the genus
<italic>Phoma</italic>. With the numerous species awaiting to be discovered, the
taxonomic system of this complex will probably be changed again as more clades
are added. Nevertheless, it is hoped that the present study on <italic>Phoma</italic>systematics, together with the “<italic>Phoma Identification
Manual</italic>”, will provide a solid foundation on which the
<italic>Didymellaceae</italic> in general, and the <italic>Phoma</italic> species in
particular, can be further delineated.</p></sec></sec></body><back><ack><p>We thank Mrs Karin Rosendahl-Peters (Plantenziektenkundige Dienst), Dr Amy
Rossman (Systematic Botany and Mycology Laboratory), Prof dr dr hc mult
Wolfgang E. Krumbein and Dr Gorbushina (University of Oldenburg) for providing
cultures. Jeroen Korving (Hubrecht Laboratory, Utrecht) is thanked for his
help in preparing the microtome sections. Dr Cecile Gueidan is kindly thanked
for her comments on <italic>Phoma paspali</italic>. Many thanks also to Mrs Trix
Merckx and Mrs Arien van Iperen who helped us with the deposit of strains and
herbarium material. Mrs Marjan Vermaas is kindly thanked for her assistance in
preparing the photoplates. This research is supported by the Dutch Ministry of
Agriculture, Nature and Food Quality through an endowment of the FES programme
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