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<record><TEI><teiHeader><fileDesc><titleStmt><title xml:lang="en">Taxonomy and phylogenetic relationships of nine species of
<italic>Hypocrea</italic> with anamorphs assignable to <italic>Trichoderma</italic> section
<italic>Hypocreanum</italic></title><author><name sortKey="Overton, Barrie E" sort="Overton, Barrie E" uniqKey="Overton B" first="Barrie E." last="Overton">Barrie E. Overton</name><affiliation><nlm:aff id="aff1"><italic>The Pennsylvania State University, Department of Plant Pathology, Buckhout Laboratory, University Park, Pennsylvania 16802, U.S.A.</italic>: Current address:<italic>Lock Haven University of Pennsylvania, Department of Biology, 119 Ulmer Hall, Lock Haven PA, 17745, U.S.A.</italic></nlm:aff></affiliation></author><author><name sortKey="Stewart, Elwin L" sort="Stewart, Elwin L" uniqKey="Stewart E" first="Elwin L." last="Stewart">Elwin L. Stewart</name><affiliation><nlm:aff id="aff2"><italic>The Pennsylvania State University, Department of Plant Pathology, Buckhout Laboratory, University Park, Pennsylvania 16802, U.S.A.</italic></nlm:aff></affiliation></author><author><name sortKey="Geiser, David M" sort="Geiser, David M" uniqKey="Geiser D" first="David M." last="Geiser">David M. Geiser</name><affiliation><nlm:aff id="aff2"><italic>The Pennsylvania State University, Department of Plant Pathology, Buckhout Laboratory, University Park, Pennsylvania 16802, U.S.A.</italic></nlm:aff></affiliation></author></titleStmt><publicationStmt><idno type="wicri:source">PMC</idno><idno type="pmid">18490989</idno><idno type="pmc">2104734</idno><idno type="url">http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2104734</idno><idno type="RBID">PMC:2104734</idno><idno type="doi">10.3114/sim.2006.56.02</idno><date when="2006">2006</date><idno type="wicri:Area/Pmc/Corpus">000102</idno><idno type="wicri:explorRef" wicri:stream="Pmc" wicri:step="Corpus" wicri:corpus="PMC">000102</idno></publicationStmt><sourceDesc><biblStruct><analytic><title xml:lang="en" level="a" type="main">Taxonomy and phylogenetic relationships of nine species of
<italic>Hypocrea</italic> with anamorphs assignable to <italic>Trichoderma</italic> section
<italic>Hypocreanum</italic></title><author><name sortKey="Overton, Barrie E" sort="Overton, Barrie E" uniqKey="Overton B" first="Barrie E." last="Overton">Barrie E. Overton</name><affiliation><nlm:aff id="aff1"><italic>The Pennsylvania State University, Department of Plant Pathology, Buckhout Laboratory, University Park, Pennsylvania 16802, U.S.A.</italic>: Current address:<italic>Lock Haven University of Pennsylvania, Department of Biology, 119 Ulmer Hall, Lock Haven PA, 17745, U.S.A.</italic></nlm:aff></affiliation></author><author><name sortKey="Stewart, Elwin L" sort="Stewart, Elwin L" uniqKey="Stewart E" first="Elwin L." last="Stewart">Elwin L. Stewart</name><affiliation><nlm:aff id="aff2"><italic>The Pennsylvania State University, Department of Plant Pathology, Buckhout Laboratory, University Park, Pennsylvania 16802, U.S.A.</italic></nlm:aff></affiliation></author><author><name sortKey="Geiser, David M" sort="Geiser, David M" uniqKey="Geiser D" first="David M." last="Geiser">David M. Geiser</name><affiliation><nlm:aff id="aff2"><italic>The Pennsylvania State University, Department of Plant Pathology, Buckhout Laboratory, University Park, Pennsylvania 16802, U.S.A.</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="2006">2006</date></imprint></series></biblStruct></sourceDesc></fileDesc><profileDesc><textClass></textClass></profileDesc></teiHeader><front><div type="abstract" xml:lang="en"><p>Morphological studies and phylogenetic analyses of DNA sequences from the
internal transcribed spacer (ITS) regions of the nuclear ribosomal gene
repeat, a partial sequence of RNA polymerase II subunit (<italic>rpb2</italic>), and a
partial sequence of the large exon of <italic>tef1</italic> (LE<italic>tef1</italic>) were
used to investigate the taxonomy and systematics of nine <italic>Hypocrea</italic>species with anamorphs assignable to <italic>Trichoderma</italic> sect.
<italic>Hypocreanum. Hypocrea corticioides</italic> and <italic>H. sulphurea</italic> are
reevaluated. Their <italic>Trichoderma</italic> anamorphs are described and the
phylogenetic positions of these species are determined. <italic>Hypocrea
sulphurea</italic> and <italic>H. subcitrina</italic> are distinct species based on
studies of the type specimens. <italic>Hypocrea egmontensis</italic> is a facultative
synonym of the older name <italic>H. subcitrina. Hypocrea</italic> with anamorphs
assignable to <italic>Trichoderma</italic> sect. <italic>Hypocreanum</italic> formed a
well-supported clade. Five species with anamorphs morphologically similar to
sect. <italic>Hypocreanum, H. avellanea, H. parmastoi, H. megalocitrina, H.
alcalifuscescens</italic>, and <italic>H. pezizoides</italic>, are not located in this
clade. <italic>Protocrea farinosa</italic> belongs to <italic>Hypocrea s.s</italic>.</p></div></front><back><div1 type="bibliography"><listBibl><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct></listBibl></div1></back></TEI><pmc article-type="research-article"><pmc-dir>properties open_access</pmc-dir>
<front><journal-meta><journal-id journal-id-type="nlm-ta">Stud Mycol</journal-id><journal-title>Studies in Mycology</journal-title><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">18490989</article-id><article-id pub-id-type="pmc">2104734</article-id><article-id pub-id-type="publisher-id">0039</article-id><article-id pub-id-type="doi">10.3114/sim.2006.56.02</article-id><article-categories><subj-group subj-group-type="heading"><subject>Articles</subject></subj-group></article-categories><title-group><article-title>Taxonomy and phylogenetic relationships of nine species of
<italic>Hypocrea</italic> with anamorphs assignable to <italic>Trichoderma</italic> section
<italic>Hypocreanum</italic></article-title></title-group><contrib-group><contrib contrib-type="author"><name><surname>Overton</surname><given-names>Barrie E.</given-names></name><xref ref-type="aff" rid="aff1">1</xref><xref ref-type="corresp" rid="cor1">*</xref></contrib><contrib contrib-type="author"><name><surname>Stewart</surname><given-names>Elwin L.</given-names></name><xref ref-type="aff" rid="aff2">2</xref></contrib><contrib contrib-type="author"><name><surname>Geiser</surname><given-names>David M.</given-names></name><xref ref-type="aff" rid="aff2">2</xref></contrib></contrib-group><aff id="aff1"><label>1</label><italic>The Pennsylvania State University, Department of Plant Pathology, Buckhout Laboratory, University Park, Pennsylvania 16802, U.S.A.</italic>: Current address:<italic>Lock Haven University of Pennsylvania, Department of Biology, 119 Ulmer Hall, Lock Haven PA, 17745, U.S.A.</italic></aff><aff id="aff2"><label>2</label><italic>The Pennsylvania State University, Department of Plant Pathology, Buckhout Laboratory, University Park, Pennsylvania 16802, U.S.A.</italic></aff><author-notes><fn id="cor1"><label>*</label><p><italic>Correspondence</italic>: Barrie E. Overton,
<email>boverton@lhup.edu</email></p></fn></author-notes><pub-date pub-type="ppub"><year>2006</year></pub-date><volume>56</volume><fpage>39</fpage><lpage>65</lpage><permissions><copyright-statement>Copyright © Copyright 2006 Centraalbureau voor Schimmelcultures, P.O.
Box 85167, 3508 AD Utrecht, The Netherlands.</copyright-statement><license><p>You are free to share–to copy, distribute and transmit the work, under the following conditions:</p><p><bold>Attribution</bold>: You must attribute the work in the manner specified by the author or licensor (but not in any way that suggests that they endorse you or your use of the work).</p><p><bold>Non-commercial</bold>: You may not use this work for commercial purposes.</p><p><bold>No derivative works</bold>: You may not alter, transform, or build upon this work.</p><p>For any reuse or distribution, you must make clear to others the license terms of this work, which can be found at <ext-link ext-link-type="uri" xlink:href="http://creativecommons.org/licenses/by-nc-nd/3.0/legalcode">http://creativecommons.org/licenses/by-nc-nd/3.0/legalcode.</ext-link> Any of the above conditions can be waived if you get permission from the copyright holder. Nothing in this license impairs or restricts the author's moral rights.</p></license></permissions><self-uri xlink:title="pdf" xlink:href="39.pdf"></self-uri><abstract><p>Morphological studies and phylogenetic analyses of DNA sequences from the
internal transcribed spacer (ITS) regions of the nuclear ribosomal gene
repeat, a partial sequence of RNA polymerase II subunit (<italic>rpb2</italic>), and a
partial sequence of the large exon of <italic>tef1</italic> (LE<italic>tef1</italic>) were
used to investigate the taxonomy and systematics of nine <italic>Hypocrea</italic>species with anamorphs assignable to <italic>Trichoderma</italic> sect.
<italic>Hypocreanum. Hypocrea corticioides</italic> and <italic>H. sulphurea</italic> are
reevaluated. Their <italic>Trichoderma</italic> anamorphs are described and the
phylogenetic positions of these species are determined. <italic>Hypocrea
sulphurea</italic> and <italic>H. subcitrina</italic> are distinct species based on
studies of the type specimens. <italic>Hypocrea egmontensis</italic> is a facultative
synonym of the older name <italic>H. subcitrina. Hypocrea</italic> with anamorphs
assignable to <italic>Trichoderma</italic> sect. <italic>Hypocreanum</italic> formed a
well-supported clade. Five species with anamorphs morphologically similar to
sect. <italic>Hypocreanum, H. avellanea, H. parmastoi, H. megalocitrina, H.
alcalifuscescens</italic>, and <italic>H. pezizoides</italic>, are not located in this
clade. <italic>Protocrea farinosa</italic> belongs to <italic>Hypocrea s.s</italic>.</p></abstract><kwd-group><kwd>Ascomycetes</kwd><kwd><italic>Hypocreales</italic></kwd><kwd><italic>Hypocreanum</italic></kwd><kwd><italic>Hypocrea corticioides</italic></kwd><kwd><italic>H. egmontensis</italic></kwd><kwd><italic>H. parmastoi</italic></kwd><kwd><italic>H. alcalifuscescens</italic></kwd><kwd><italic>H. subsulphurea</italic></kwd><kwd><italic>H. farinosa</italic></kwd><kwd><italic>H. subcitrina</italic></kwd><kwd><italic>H. sulphurea</italic></kwd><kwd><italic>H. victoriensis</italic></kwd><kwd>ITS rDNA</kwd><kwd><italic>Lentinula edodes</italic></kwd><kwd>systematics</kwd><kwd><italic>rpb2</italic> gene sequences</kwd><kwd><italic>tef1</italic> gene sequences</kwd><kwd><italic>Trichoderma</italic></kwd></kwd-group></article-meta><notes><fn-group><fn><p><bold>Taxonomic novelties:</bold> <italic>Hypocrea eucorticioides</italic> Overton, nom.
nov., <italic>Hypocrea victoriensis</italic> Overton, sp. nov.<italic>, Hypocrea
parmastoi</italic> Overton, sp. nov., <italic>Hypocrea alcalifuscescens</italic> Overton,
sp. nov.</p></fn></fn-group></notes></front><body><sec><title>INTRODUCTION</title><p>Nine species of <italic>Hypocrea</italic> Fr. (<italic>Ascomycetes, Hypocreales,
Hypocreaceae</italic>) with effused stromata from Japan, Australia, New Zealand,
North America, Europe, and Central America, are newly described or
redescribed. Anamorphs of these species are morphologically similar, having
acremonium- or verticillium-like conidiophores with hyaline conidia, and are
assignable to <italic>Trichoderma</italic> sect<italic>. Hypocreanum</italic> Bissett.</p><p><italic>Hypocrea sulphurea</italic> (Schw.) Sacc. is a common, yellow, effused
fungicolous species recorded from North America and Europe that occurs on
<italic>Exidia</italic> spp. Dingley
(<xref ref-type="bibr" rid="ref2">1956</xref>) considered <italic>H.
subcitrina</italic> Kalchbr. & Cooke, recorded from Africa, as a synonym of
the older <italic>H. sulphurea</italic>, but this synonymy has never been critically
examined. Dingley (<xref ref-type="bibr" rid="ref2">1956</xref>)
published the new name <italic>H. egmontensis</italic> from New Zealand based on a
fungus with yellow effused stromata. The relationship between <italic>H.
egmontensis</italic> and <italic>H. sulphurea</italic> has not been established. Doi
(<xref ref-type="bibr" rid="ref5">1972</xref>) described <italic>Hypocrea
sulphurea</italic> f. <italic>macrospora</italic> Yoshim. Doi. We compared morphologically
type material (NY) of this forma with collections of <italic>H. sulphurea</italic>from North America and Europe. A specimen identified as <italic>Hypocrea
subsulphurea</italic> Syd. in De Wild. was recently collected and cultured in
Japan and redescribed. <italic>Hypocrea corticioides</italic> Speg. is similar in
appearance to <italic>H. sulphurea</italic>, but <italic>H. corticioides</italic> occurs on
decorticated wood and has a tropical distribution. <italic>Hypocrea
corticioides</italic> Speg. is a later homonym of <italic>H. corticioides</italic> Berk.
& Broome. The type material of <italic>H. corticioides</italic> Berk. & Broome
is indistinguishable from and therefore synonymous with <italic>Stilbocrea
macrostoma</italic> (Berk. & M.A. Curtis) Höhn., a member of the
<italic>Bionectriaceae</italic> (<xref ref-type="bibr" rid="ref15">Rossman <italic>et
al.</italic> 1999</xref>). A new name is proposed for <italic>H.
corticioides</italic> Speg.</p><p>Two apparently new species of <italic>Hypocrea</italic> with hyphal stromata were
studied. Their relationship to <italic>Hypocrea</italic> spp. with
pseudoparenchymatous tissue was unclear. In addition, the relationship of
these hyphal species to <italic>Protocrea farinosa</italic> (Berk. & Broome)
Petch, which also has a hyphal stroma, had to be examined.</p><p>Kullnig-Gradinger <italic>et al.</italic>(<xref ref-type="bibr" rid="ref11">2002</xref>) showed that some
<italic>Trichoderma</italic> species with anamorphs in <italic>Trichoderma</italic> sect.
<italic>Hypocreanum</italic> form a highly supported subclade of sect. <italic>Pachybasium
sensu lato</italic> and suggested that sections <italic>Hypocreanum</italic> and
<italic>Pachybasium</italic> are phylogenetically indistinguishable. Their analysis
included a limited number of taxa with acremonium- or verticillium-like
anamorphs. More recently, Chaverri <italic>et al.</italic>(<xref ref-type="bibr" rid="ref1">2003</xref>) used partial sequences
of the RNA polymerase II subunit (<italic>rpb2</italic>) and the large exon of
tef-1α (LE<italic>tef1</italic>) and found that anamorphs referable to sect.
<italic>Hypocreanum</italic> do not form a monophyletic group, as <italic>H.
pezizoides</italic> Berk. & Broome and <italic>H. avellanea</italic> S.T. Carey &
Rogerson were situated in the <italic>H. rufa</italic> clade. Chaverri <italic>et al.</italic>(<xref ref-type="bibr" rid="ref1">2003</xref>) showed that <italic>H.
citrina</italic> (Pers.: Fr.) Fr. and <italic>H. pulvinata</italic> Fuckel form a highly
supported clade, the limits of which were not established. Dodd <italic>et
al.</italic> (<xref ref-type="bibr" rid="ref3">2002</xref>) showed, using
the ITS1-5.8S-ITS2 rDNA (ITS) region, that <italic>H. pulvinata</italic> and <italic>H.
sulphurea</italic> form two distinct subclades of a strongly supported but
phylogenetically unresolved clade. These authors did not conclude that sect.
<italic>Hypocreanum</italic> and sect. <italic>Pachybasium</italic> were phylogenetically
indistinguishable. The results of Dodd <italic>et al.</italic>(<xref ref-type="bibr" rid="ref3">2002</xref>) and Chaverri <italic>et
al.</italic> (<xref ref-type="bibr" rid="ref1">2003</xref>) support the
conclusion of Kullnig-Gradinger <italic>et al.</italic>(<xref ref-type="bibr" rid="ref11">2002</xref>) that section
<italic>Pachybasium</italic> is paraphyletic. The seven species included are compared
to selected species treated by Overton <italic>et al.</italic>(<xref ref-type="bibr" rid="ref12">2006</xref>) to establish the
phylogenetic limits of <italic>Trichoderma</italic> sect. <italic>Hypocreanum</italic>.</p><p>The objectives of this study are: (1) to determine whether <italic>H.
sulphurea, H. subcitrina</italic>, and <italic>H. egmontensis</italic> are distinct
species; (2) to verify the phylogenetic relationship between <italic>H.
subsulphurea</italic> and <italic>H. sulphurea</italic>; (3) to verify the relationship
between <italic>H. corticioides</italic> and <italic>H. sulphurea</italic>; (4) to determine
the relationships of two new hyphal species to <italic>Protocrea farinosa</italic>;
(5) to investigate the phylogenetic boundaries of <italic>Hypocrea</italic> with
anamorphs in <italic>Trichoderma</italic> sect. <italic>Hypocreanum</italic>; and (6) to
describe the phylogenetic species delineated in this study according to
criteria developed by Taylor <italic>et al.</italic>(<xref ref-type="bibr" rid="ref20">2000</xref>).</p></sec><sec sec-type="materials|methods"><title>MATERIALS AND METHODS</title><sec><title>Collections and isolates</title><p>Doi's illustrations and descriptions
(<xref ref-type="bibr" rid="ref4">1971</xref>,
<xref ref-type="bibr" rid="ref5">1972</xref>,
<xref ref-type="bibr" rid="ref6">1975</xref>) were used in making
initial species determinations. <xref ref-type="table" rid="tbl1">Table
1</xref> lists the accession numbers used in this study. Frequently cited
collectors are abbreviated: B.E. Overton (B.E.O.), G. J. Samuels (G.J.S.), and
K. Põldmaa (K.P.). All isolates with G.J.S. designations were obtained
by isolating single ascospores on CMD with the aid of a micromanipulator. All
isolates with B.E.O. designations were obtained from plating the entire
contents of individual perithecia. Unless otherwise noted, host and substratum
data are taken from herbarium labels. The presentation of measurements is the
same as in Overton <italic>et al.</italic>(<xref ref-type="bibr" rid="ref12">2006</xref>).</p><p><table-wrap position="float" id="tbl1"><label>Table 1.</label><caption><p>Isolates used in molecular phylogenetic analyses. (* = ex-type strain).</p></caption><table frame="hsides" rules="groups"><thead><tr><th colspan="1" rowspan="1" align="left" valign="top"><bold>Name</bold></th><th colspan="1" rowspan="1" align="left" valign="top"><bold>Accession number</bold></th><th colspan="1" rowspan="1" align="left" valign="top"><bold>Origin</bold></th><th colspan="3" rowspan="1" align="center" valign="bottom"><bold>GenBank accession number</bold><hr></hr></th></tr><tr><th colspan="1" rowspan="1" align="left" valign="top"></th><th colspan="1" rowspan="1" align="left" valign="top"></th><th colspan="1" rowspan="1" align="left" valign="top"></th><th colspan="1" rowspan="1" align="center" valign="top"><bold>ITS</bold></th><th colspan="1" rowspan="1" align="center" valign="top"><bold>LE<italic>tef1</italic></bold></th><th colspan="1" rowspan="1" align="center" valign="top"><bold><italic>rpb2</italic></bold></th></tr></thead><tbody><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>H. pulvinata</italic> Fuckel
</td><td colspan="1" rowspan="1" align="left" valign="top"> G.J.S. 94-20
</td><td colspan="1" rowspan="1" align="left" valign="top"> Tushar Mountains, Utah, U.S.A.
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835409
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835485
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835451
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"></td><td colspan="1" rowspan="1" align="left" valign="top"> G.J.S. 92-127
</td><td colspan="1" rowspan="1" align="left" valign="top"> Olympia National Park, Washington, U.S.A.
</td><td colspan="1" rowspan="1" align="center" valign="top"> AF487666
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835484
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835461
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"></td><td colspan="1" rowspan="1" align="left" valign="top"> G.J.S. 98-104
</td><td colspan="1" rowspan="1" align="left" valign="top"> Naturpark Saar-Hundsrück, Germany
</td><td colspan="1" rowspan="1" align="center" valign="top"> AF487665
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835490
</td><td colspan="1" rowspan="1" align="center" valign="top"> AF545559
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"></td><td colspan="1" rowspan="1" align="left" valign="top"> G.J.S. 95-220
</td><td colspan="1" rowspan="1" align="left" valign="top"> Waldviertel, Lower Austria, Austria
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835407
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835486
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835452
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>H. americana</italic> (Canham) Overton
</td><td colspan="1" rowspan="1" align="left" valign="top"> G.J.S. 92-93
</td><td colspan="1" rowspan="1" align="left" valign="top"> New Mexico, U.S.A.
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835410
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835489
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835455
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"></td><td colspan="1" rowspan="1" align="left" valign="top"> G.J.S. 94-79
</td><td colspan="1" rowspan="1" align="left" valign="top"> White Mountains, Arizona, U.S.A.
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835408
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835491
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835456
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>H. protopulvinata</italic> Yoshim. Doi
</td><td colspan="1" rowspan="1" align="left" valign="top"> K.P. 00-56
</td><td colspan="1" rowspan="1" align="left" valign="top"> Unknown, U.S.A.
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835406
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835488
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835453
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"></td><td colspan="1" rowspan="1" align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=739.83&link_type=CBS">CBS 739.83*</ext-link></td><td colspan="1" rowspan="1" align="left" valign="top"> Chiba Pref., Kiyosumi, Fudagou, Japan
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835405
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835487
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835463
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>H. citrina</italic> (Pers. : Fr.) <italic>Fr.</italic></td><td colspan="1" rowspan="1" align="left" valign="top"> G.J.S. 95-183
</td><td colspan="1" rowspan="1" align="left" valign="top"> Daniel Boone National Forest, Kentucky, U.S.A.
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835413
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835469
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835458
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"></td><td colspan="1" rowspan="1" align="left" valign="top"> B.E.O. 99-29
</td><td colspan="1" rowspan="1" align="left" valign="top"> Oswego County, New York, U.S.A.
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835412
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835482
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835464
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"></td><td colspan="1" rowspan="1" align="left" valign="top"> G.J.S. 89-145
</td><td colspan="1" rowspan="1" align="left" valign="top"> Devon, Budleigh, Saltaton, U.K.
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835414
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835483
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835457
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"></td><td colspan="1" rowspan="1" align="left" valign="top"> G.J.S. 96-275
</td><td colspan="1" rowspan="1" align="left" valign="top"> Ascutung, Vermont, U.S.A.
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835418
</td><td colspan="1" rowspan="1" align="right" valign="top"> -
</td><td colspan="1" rowspan="1" align="right" valign="top"> -
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"></td><td colspan="1" rowspan="1" align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=708.73&link_type=CBS">CBS 708.73</ext-link></td><td colspan="1" rowspan="1" align="left" valign="top"> Baam, Zandheuvelweg, Netherlands
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835415
</td><td colspan="1" rowspan="1" align="right" valign="top"> -
</td><td colspan="1" rowspan="1" align="right" valign="top"> -
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"></td><td colspan="1" rowspan="1" align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=853.70&link_type=CBS">CBS 853.70</ext-link></td><td colspan="1" rowspan="1" align="left" valign="top"> Pelmer Wald near Gerolstein, Germany
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835416
</td><td colspan="1" rowspan="1" align="right" valign="top"> -
</td><td colspan="1" rowspan="1" align="right" valign="top"> -
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"></td><td colspan="1" rowspan="1" align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=894.85&link_type=CBS">CBS 894.85*</ext-link></td><td colspan="1" rowspan="1" align="left" valign="top"> Hestreux near Eupen, Belgium
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835417
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835481
</td><td colspan="1" rowspan="1" align="center" valign="top"> AF545561
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>H. pseudostraminea</italic> Yoshim. Doi
</td><td colspan="1" rowspan="1" align="left" valign="top"> G.J.S. 91-135
</td><td colspan="1" rowspan="1" align="left" valign="top"> Prince Georges County, Maryland, U.S.A.
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835419
</td><td colspan="1" rowspan="1" align="right" valign="top"> -
</td><td colspan="1" rowspan="1" align="right" valign="top"> -
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"></td><td colspan="1" rowspan="1" align="left" valign="top"> G.J.S. 90-74
</td><td colspan="1" rowspan="1" align="left" valign="top"> Dutches County, New York, U.S.A.
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835420
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835470
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835454
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"></td><td colspan="1" rowspan="1" align="left" valign="top"> G.J.S. 95-169
</td><td colspan="1" rowspan="1" align="left" valign="top"> Daniel Boon National Forest, Kentucky, U.S.A.
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835421
</td><td colspan="1" rowspan="1" align="right" valign="top"> -
</td><td colspan="1" rowspan="1" align="right" valign="top"> -
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"></td><td colspan="1" rowspan="1" align="left" valign="top"> G.J.S. 95-189
</td><td colspan="1" rowspan="1" align="left" valign="top"> Brown County, Indiana, U.S.A.
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835422
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835480
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835459
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"></td><td colspan="1" rowspan="1" align="left" valign="top"> B.E.O. 99-36
</td><td colspan="1" rowspan="1" align="left" valign="top"> Patapsco State Park, Maryland, U.S.A.
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835423
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835468
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835465
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>H. megalocitrina</italic> Yoshim. Doi
</td><td colspan="1" rowspan="1" align="left" valign="top"> B.E.O. 00-09
</td><td colspan="1" rowspan="1" align="left" valign="top"> North Carolina, U.S.A.
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835511
</td><td colspan="1" rowspan="1" align="center" valign="top"> AY225855
</td><td colspan="1" rowspan="1" align="center" valign="top"> AF545563
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>H. microcitrina</italic> Yoshim.Doi
</td><td colspan="1" rowspan="1" align="left" valign="top"> G.J.S. 97-248
</td><td colspan="1" rowspan="1" align="left" valign="top"> Georgia, U.S.A.
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835424
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835479
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835462
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"></td><td colspan="1" rowspan="1" align="left" valign="top"> G.J.S. 91-61
</td><td colspan="1" rowspan="1" align="left" valign="top"> Virginia, U.S.A.
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835426
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835478
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835460
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>H. sulphurea</italic> (Schw.) Sacc.
</td><td colspan="1" rowspan="1" align="left" valign="top"> G.J.S. 95-190
</td><td colspan="1" rowspan="1" align="left" valign="top"> Indiana, U.S.A.
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835425
</td><td colspan="1" rowspan="1" align="center" valign="top"> AY225858
</td><td colspan="1" rowspan="1" align="center" valign="top"> AF545560
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"></td><td colspan="1" rowspan="1" align="left" valign="top"> G.J.S. 95-140
</td><td colspan="1" rowspan="1" align="left" valign="top"> Styria, Austria
</td><td colspan="1" rowspan="1" align="center" valign="top"> AF487664
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835471
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835515
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"></td><td colspan="1" rowspan="1" align="left" valign="top"> G.J.S. 00-172
</td><td colspan="1" rowspan="1" align="left" valign="top"> Moscow, Russia
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835510
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835493
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835523
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>H. victoriensis</italic> Overton
</td><td colspan="1" rowspan="1" align="left" valign="top"> G.J.S. 99-130
</td><td colspan="1" rowspan="1" align="left" valign="top"> Victoria, Australia
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835504
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835472
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835516
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"></td><td colspan="1" rowspan="1" align="left" valign="top"> G.J.S. 99-200*
</td><td colspan="1" rowspan="1" align="left" valign="top"> Victoria, Australia
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835505
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835473
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835517
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"></td><td colspan="1" rowspan="1" align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=500.67&link_type=CBS">CBS 500.67</ext-link></td><td colspan="1" rowspan="1" align="left" valign="top"> New Zealand
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835466
</td><td colspan="1" rowspan="1" align="right" valign="top"> -
</td><td colspan="1" rowspan="1" align="right" valign="top"> -
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>H. eucorticioides</italic> Overton
</td><td colspan="1" rowspan="1" align="left" valign="top"> G.J.S. 99-61
</td><td colspan="1" rowspan="1" align="left" valign="top"> Limon, Costa Rica
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835467
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835474
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835518
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>H. subsulphurea</italic> Kalchbr. & Cooke
</td><td colspan="1" rowspan="1" align="left" valign="top"> M 141
</td><td colspan="1" rowspan="1" align="left" valign="top"> Kurokami Kumamoto, Japan
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835509
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835492
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835522
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>H. farinosa</italic> Berk. & Broome
</td><td colspan="1" rowspan="1" align="left" valign="top"> G.J.S. 91-101
</td><td colspan="1" rowspan="1" align="left" valign="top"> Maryland, U.S.A.
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835507
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835476
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835520
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"></td><td colspan="1" rowspan="1" align="left" valign="top"> G.J.S. 89-139
</td><td colspan="1" rowspan="1" align="left" valign="top"> Unknown mushroom farm, U.S.A.
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835508
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835477
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835521
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>H. alcalifuscescens</italic> Overton
</td><td colspan="1" rowspan="1" align="left" valign="top"> B.E.O. 99-16
</td><td colspan="1" rowspan="1" align="left" valign="top"> Maryland, U.S.A.
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835506
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835475
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835519
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"></td><td colspan="1" rowspan="1" align="left" valign="top"> TFC 181548*
</td><td colspan="1" rowspan="1" align="left" valign="top"> Estonia
</td><td colspan="1" rowspan="1" align="right" valign="top"> -
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ834455
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ834462
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>H. parmastoi</italic> Overton
</td><td colspan="1" rowspan="1" align="left" valign="top"> TFC 97-143*
</td><td colspan="1" rowspan="1" align="left" valign="top"> Voru Commune, Estonia
</td><td colspan="1" rowspan="1" align="right" valign="top"> -
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ834456
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ834463
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>H. avellanea</italic> Carey & Rogerson
</td><td colspan="1" rowspan="1" align="left" valign="top"> C.T.R. 77-155*
</td><td colspan="1" rowspan="1" align="left" valign="top"> Type locality, Massachusetts, U.S.A.
</td><td colspan="1" rowspan="1" align="right" valign="top"> -
</td><td colspan="1" rowspan="1" align="center" valign="top"> AY225857
</td><td colspan="1" rowspan="1" align="center" valign="top"> AF545562
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>H.</italic> cf. <italic>ochroleuca</italic></td><td colspan="1" rowspan="1" align="left" valign="top"> G.J.S. 01-265
</td><td colspan="1" rowspan="1" align="left" valign="top"> Thailand
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835512
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835494
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835524
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>H. pezizoides</italic> Berk. & Broome
</td><td colspan="1" rowspan="1" align="left" valign="top"> G.J.S. 01-231
</td><td colspan="1" rowspan="1" align="left" valign="top"> Unknown
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835513
</td><td colspan="1" rowspan="1" align="center" valign="top"> AY225859
</td><td colspan="1" rowspan="1" align="center" valign="top"> AF545564
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>H.</italic> cf. <italic>cinereoflava</italic></td><td colspan="1" rowspan="1" align="left" valign="top"> G.J.S. 92-102
</td><td colspan="1" rowspan="1" align="left" valign="top"> Unknown
</td><td colspan="1" rowspan="1" align="right" valign="top"> -
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ834454
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ834461
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>H. psychrophila</italic> E. Müll., Aebi & J. Webster
</td><td colspan="1" rowspan="1" align="left" valign="top"> HY8 (<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=262.71&link_type=CBS">CBS 262.71</ext-link>)
</td><td colspan="1" rowspan="1" align="left" valign="top"> Switzerland
</td><td colspan="1" rowspan="1" align="right" valign="top"> -
</td><td colspan="1" rowspan="1" align="center" valign="top"> AF534584
</td><td colspan="1" rowspan="1" align="center" valign="top"> AF545520
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>H. rufa</italic> (Pers.) Fr.
</td><td colspan="1" rowspan="1" align="left" valign="top"> G.J.S. 89-127
</td><td colspan="1" rowspan="1" align="left" valign="top"> North Carolina, U.S.A.
</td><td colspan="1" rowspan="1" align="right" valign="top"> -
</td><td colspan="1" rowspan="1" align="center" valign="top"> AF534585
</td><td colspan="1" rowspan="1" align="center" valign="top"> AF545521
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>H. pilulifera</italic> J. Webster & Rifai
</td><td colspan="1" rowspan="1" align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=814.68&link_type=CBS">CBS 814.68</ext-link></td><td colspan="1" rowspan="1" align="left" valign="top"> Yorkshire, U.K.
</td><td colspan="1" rowspan="1" align="right" valign="top"> -
</td><td colspan="1" rowspan="1" align="center" valign="top"> AF534583
</td><td colspan="1" rowspan="1" align="center" valign="top"> AF545519
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>H. lutea</italic> (Tode) Petch
</td><td colspan="1" rowspan="1" align="left" valign="top"> G.J.S. 89-129
</td><td colspan="1" rowspan="1" align="left" valign="top"> New York, U.S.A
</td><td colspan="1" rowspan="1" align="right" valign="top"> -
</td><td colspan="1" rowspan="1" align="center" valign="top"> AF534581
</td><td colspan="1" rowspan="1" align="center" valign="top"> AF545517
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>H. strictipilosa</italic> Chaverri & Samuels
</td><td colspan="1" rowspan="1" align="left" valign="top"> G.J.S. 89-115
</td><td colspan="1" rowspan="1" align="left" valign="top"> Maryland, U.S.A.
</td><td colspan="1" rowspan="1" align="right" valign="top"> -
</td><td colspan="1" rowspan="1" align="center" valign="top"> AF534596
</td><td colspan="1" rowspan="1" align="center" valign="top"> AF545528
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>H. aureoviridis</italic> Plowr. & Cooke
</td><td colspan="1" rowspan="1" align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=245.63&link_type=CBS">CBS 245.63</ext-link></td><td colspan="1" rowspan="1" align="left" valign="top"> England, U.S.A.
</td><td colspan="1" rowspan="1" align="right" valign="top"> -
</td><td colspan="1" rowspan="1" align="center" valign="top"> AF534575
</td><td colspan="1" rowspan="1" align="center" valign="top"> AF545504
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>H. nigrovirens</italic> Chaverri & Samuels
</td><td colspan="1" rowspan="1" align="left" valign="top"> G.J.S. 99-64
</td><td colspan="1" rowspan="1" align="left" valign="top"> Limón, Costa Rica
</td><td colspan="1" rowspan="1" align="right" valign="top"> -
</td><td colspan="1" rowspan="1" align="center" valign="top"> AF534582
</td><td colspan="1" rowspan="1" align="center" valign="top"> AF545518
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>Trichoderma</italic> cf. <italic>citrinoviride</italic></td><td colspan="1" rowspan="1" align="left" valign="top"> G.J.S. 01-364
</td><td colspan="1" rowspan="1" align="left" valign="top"> Unknown
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835514
</td><td colspan="1" rowspan="1" align="center" valign="top"> AY225860
</td><td colspan="1" rowspan="1" align="center" valign="top"> AF545565
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>T. fertile</italic> Bissett
</td><td colspan="1" rowspan="1" align="left" valign="top"> DAOM 167070
</td><td colspan="1" rowspan="1" align="left" valign="top"> Canada
</td><td colspan="1" rowspan="1" align="right" valign="top"> -
</td><td colspan="1" rowspan="1" align="center" valign="top"> AF534617
</td><td colspan="1" rowspan="1" align="center" valign="top"> AF545545
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>T. aggressivum</italic> Samuels & W. Gams
</td><td colspan="1" rowspan="1" align="left" valign="top"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=100525&link_type=CBS">CBS 100525</ext-link></td><td colspan="1" rowspan="1" align="left" valign="top"> United Kingdom
</td><td colspan="1" rowspan="1" align="right" valign="top"> -
</td><td colspan="1" rowspan="1" align="center" valign="top"> AF534614
</td><td colspan="1" rowspan="1" align="center" valign="top"> AF545541
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>T. flavofuscum</italic> (J.M. Mill., Giddens, A.A. Foster) Bissett
</td><td colspan="1" rowspan="1" align="left" valign="top"> DAOM 167652
</td><td colspan="1" rowspan="1" align="left" valign="top"> Georgia, U.S.A.
</td><td colspan="1" rowspan="1" align="right" valign="top"> -
</td><td colspan="1" rowspan="1" align="center" valign="top"> AF534619
</td><td colspan="1" rowspan="1" align="center" valign="top"> AF545547
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>T. stromaticum</italic> Samuels & Pardo-Schultheiss
</td><td colspan="1" rowspan="1" align="left" valign="top"> P.C. 209
</td><td colspan="1" rowspan="1" align="left" valign="top"> Brazil
</td><td colspan="1" rowspan="1" align="right" valign="top"> -
</td><td colspan="1" rowspan="1" align="center" valign="top"> AF534613
</td><td colspan="1" rowspan="1" align="center" valign="top"> AF545539
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>Arachnocrea scabrida</italic> Yoshim. Doi
</td><td colspan="1" rowspan="1" align="left" valign="top"> B.E.O. 02-01
</td><td colspan="1" rowspan="1" align="left" valign="top"> New York, U.S.A.
</td><td colspan="1" rowspan="1" align="right" valign="top"> -
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ834457
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ834458
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>Sphaerostilbella</italic> cf. <italic>aureonitens</italic></td><td colspan="1" rowspan="1" align="left" valign="top"> G.J.S. 74-87
</td><td colspan="1" rowspan="1" align="left" valign="top"> New Zealand
</td><td colspan="1" rowspan="1" align="right" valign="top"> -
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ834452
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ834460
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>S.</italic> cf. <italic>aureonitens</italic></td><td colspan="1" rowspan="1" align="left" valign="top">G.J.S. 82-40
</td><td colspan="1" rowspan="1" align="left" valign="top">New Zealand
</td><td colspan="1" rowspan="1" align="right" valign="top">-
</td><td colspan="1" rowspan="1" align="center" valign="top">DQ834453
</td><td colspan="1" rowspan="1" align="center" valign="top">DQ834459
</td></tr></tbody></table></table-wrap></p></sec><sec><title>Molecular phylogenetic analyses</title><p>DNA sequence analysis was conducted using three gene sequences: ITS
1-5.8S-ITS2 (ITS), a partial sequence of the large exon of translation
elongation factor (LE<italic>tef1</italic>), and a partial sequence of the RNA
polymerase II subunit (<italic>rpb2</italic>). ITS and <italic>rpb2</italic> sequences were
generated following the protocol and primers described in Overton <italic>et
al</italic>. (<xref ref-type="bibr" rid="ref12">2006</xref>). The following
primers were employed for amplifying the LE<italic>tef1</italic> regions which differs
from the <italic>tef1</italic>region amplified in Overton <italic>et al</italic>.
(<xref ref-type="bibr" rid="ref12">2006</xref>): for LE<italic>tef1</italic>,
EF1-983F (5'-GC(C/T)CC(C/T)GG(A/C/T)CA(C/T)GGTGA(C/T)TT(C/T)AT-3') (Carbone
& Kohn 1999), EF1-2218R (5'-ATGAC(A/G)TG(A/G)GC(A/G)AC(A/G)GT(C/T)TG-3')
(S.A. Rehner, pers. comm.). Two percent dimethyl sulfoxide (DMSO) from
AMRESCO® was added to each 50 μL PCR reaction. PCR products were
purified and sequenced following the protocol in Overton <italic>et al.</italic>(<xref ref-type="bibr" rid="ref12">2006</xref>). Sequences were
assembled using SeqMan® II option and aligned using Clustal W in DNA Star
(DNA Star Inc., Madison, Wisconsin), and a phylogenetic analysis was performed
using PAUP* v. 4.0 b4 (<xref ref-type="bibr" rid="ref19">Swofford
1999</xref>). Alignments were manually adjusted in PAUP*. Outgroup taxa
varied depending on the phylogenetic analysis to meet two different objectives
in this study. For the first objective, ITS, <italic>rpb2</italic>, and
LE<italic>tef1</italic> were evaluated in single and combined analyses to establish
phylogenetic species limits. These analyses excluded the taxa <italic>H.
avellanea, H. parmastoi, H. cinereoflava</italic> Samuels & Seifert, and
<italic>H. alcalifuscescens,</italic> with isolates of <italic>T.</italic> cf.
<italic>citrinoviride, H. megalocitrina, H. pezizoides</italic>, and <italic>H.</italic> cf.
<italic>ochroleuca</italic> used as outgroup taxa. The second objective was to place
<italic>Hypocrea</italic> isolates with <italic>Trichoderma</italic> sect.
<italic>Hypocreanum</italic> anamorphs in phylogenetic context with other
<italic>Hypocrea</italic>/<italic>Trichoderma</italic> species. For the second objective,
<italic>Sphaerostilbella</italic> cf. <italic>aureonitens, Arachnocrea scabrida</italic>Yoshim. Doi., and <italic>Hypomyces stephanomatis</italic> Rogerson & Samuels were
used as outgroup taxa for the combined LE<italic>tef1</italic> and <italic>rpb2</italic>analysis with representative isolates from the different sections of
<italic>Trichoderma</italic> included in the analysis. Maximum parsimony (MP) analyses
were done using the heuristic search option under the following conditions:
TBR branch swapping, 10 random addition sequences, and gaps
(insertions/deletions) treated as missing. Bootstrap analysis was performed in
500 replicates with random sequence addition (10 replicates). For the combined
LE<italic>tef1</italic> and <italic>rpb2</italic> analysis, sequences were trimmed to the same
starting position because some GenBank sequences not generated in this study
were significantly shorter. All sequences and alignments were deposited in
GenBank (<xref ref-type="table" rid="tbl1">Table 1</xref>).</p><p>Alternate phylogenetic hypotheses reflecting different species
relationships were compared by the Kishino-Hasegawa (K-H) test
(<xref ref-type="table" rid="tbl2">Table 2</xref>) in PAUP* for the
combined LE<italic>tef1</italic> and <italic>rpb2</italic> data set. The most parsimonious
trees recovered with and without constraints were compared by likelihood
scores (<xref ref-type="table" rid="tbl2">Table 2</xref>). The
likelihood model implemented in the K-H test assumed equal rates of
substitution and empirical base frequencies. Models of sequence evolution were
tested and model parameters obtained for the LE<italic>tef1, rpb2</italic>, and
combined alignments using MODELTEST 3.06
(<xref ref-type="bibr" rid="ref14">Posada & Crandall 1998</xref>)
as implemented in PAUP*. For the LE<italic>tef1</italic> data, the likelihood ratio
test (LRT) implemented in MODELTEST, selected the TIM+I+G model with unequal
base frequencies; nucleotide frequencies were set to A: 0.2133, C: 0.3337, G:
0.2211, T: 0.2320; a gamma-shape parameter of 0.5234; and substitution rates
set to 1.0000 (A–C), 3.1252 (A–G), 1.6847 (A–T), 1.6847
(C–G), 10.5209 (C–T), and 1.0000 (G–T). For the
<italic>rpb2</italic> data, the LRT implemented in MODELTEST, selected the TrN+I+G
model with unequal base frequencies; nucleotide frequencies were set to A:
0.2413, C: 0.2787, G: 0.2551, T: 0.2248; a gamma-shape parameter of 1.1736;
and substitution rates set to 1.0000 (A–C), 6.5499 (A–G), 1.0000
(A–T), 1.0000 (C–G); 9.0762 (C–T), and 1.0000 (G–T).
For the combined LE<italic>tef1</italic> and <italic>rpb2</italic> data set, the LRT
implemented in MODELTEST, selected GTR G+I model with unequal base
frequencies; nucleotide frequencies were set to A: 0.22590, C: 0.30330, G:
0.24090, T: 0.22990; a gamma-shape parameter of 0.87796; and substitution
rates set to 1.0000 (A–C), 5.2773 (A–G), 1.0000 (A–T),
1.0000 (C–G), 8.4309 (C–T), and 1.0000 (G–T). A maximum
likelihood (ML) tree was then obtained in PAUP* using 10 random sequence
addition replicates and the substitution model suggested by MODELTEST.
Bootstrap analysis was performed with 500 replicates and fast stepwise
addition.</p><p><table-wrap position="float" id="tbl2"><label>Table 2.</label><caption><p>Results of the Kishino-Hasegawa likelihood test</p></caption><table frame="hsides" rules="groups"><thead><tr><th colspan="1" rowspan="1" align="left" valign="top"><bold>Topology</bold></th><th colspan="1" rowspan="1" align="left" valign="top"><bold>Trees</bold></th><th colspan="1" rowspan="1" align="left" valign="top"><bold>—In likelihood</bold></th><th colspan="1" rowspan="1" align="left" valign="top"><bold>P</bold><xref ref-type="table-fn" rid="tblfn1"><bold><sup>1</sup></bold></xref></th></tr></thead><tbody><tr><td colspan="1" rowspan="1" align="left" valign="top"> Unconstrained
</td><td colspan="1" rowspan="1" align="left" valign="top"> 1<xref ref-type="table-fn" rid="tblfn2">2</xref></td><td colspan="1" rowspan="1" align="center" valign="top"> 14817.41
</td><td colspan="1" rowspan="1" align="center" valign="top"> Best
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"> Monophyletic <italic>Hypocreanum</italic></td><td colspan="1" rowspan="1" align="left" valign="top"> 5
</td><td colspan="1" rowspan="1" align="center" valign="top"> 15048.56-15048.67
</td><td colspan="1" rowspan="1" align="center" valign="top"> <0.0001*
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top">Monophyletic Hyphal
</td><td colspan="1" rowspan="1" align="left" valign="top">1
</td><td colspan="1" rowspan="1" align="center" valign="top">15156.58
</td><td colspan="1" rowspan="1" align="center" valign="top"><0.0001*
</td></tr></tbody></table><table-wrap-foot><fn id="tblfn1"><label>1</label><p>Probability of getting a more extreme T-value under the null hypothesis of
no difference between the two trees (two-tailed test); indicates significance
at P < 0.05.</p></fn><fn id="tblfn2"><label>2</label><p>The best —In likelihood tree from the maximum parsimony analysis.</p></fn></table-wrap-foot></table-wrap></p></sec><sec><title>Morphology</title><p>Anamorph and teleomorph characteristics were measured from isolates and
specimens representative of each phylogenetic species. Cultures of
<italic>Hypocrea</italic> were grown on PDA, CMD and SNA at 20°C, with 12 h
fluorescent light and 12 h darkness. Observations of anamorphs were made at
<italic>ca.</italic> 7–10 d post inoculation. Anamorph and teleomorph characters
were measured following Overton <italic>et al</italic>.
(<xref ref-type="bibr" rid="ref12">2006</xref>) with the exception that
optimal growth temperatures were not determined. Colour terminology was
obtained from Kornerup & Wanscher
(<xref ref-type="bibr" rid="ref8">1981</xref>). Important morphological
characters used in species recognition are discussed in the comments section
immediately following each species description.</p><p><fig position="float" id="fig1"><label>Fig. 1.</label><caption><p>Parsimony analysis of ITS. One phylogram of 3193 most parsimonious trees;
217 steps; consistency index 0.779; retention index 0.855; homoplasy index
0.221; numerical values of branch lengths are given above and bootstrap values
(500 replicates with 10 random addition replications) are indicated below
branches. Outgroup taxa: <italic>H. megalocitrina</italic>; <italic>H. ochroleuca</italic>;
<italic>H. pezizoides</italic>; <italic>T.</italic> cf<italic>. citrinoviride</italic>.</p></caption><graphic xlink:href="39fig1"></graphic></fig></p></sec></sec><sec><title>RESULTS</title><sec><title>Phylogeny</title><p>Except for minor differences, the gene trees are concordant (Figs
<xref rid="fig1" ref-type="fig">1</xref>,
<xref rid="fig2" ref-type="fig">2</xref>,
<xref rid="fig3" ref-type="fig">3</xref>). The gene tree generated
from ITS is slightly different from those obtained from LE<italic>tef1</italic> and
<italic>rpb2. Hypocrea sulphurea</italic> isolate G.J.S 00-172 from Russia grouped
with North American isolates in the ITS tree
(<xref rid="fig1" ref-type="fig">Fig. 1</xref>) but grouped with
G.J.S. 95-140 from Europe in <italic>rpb2</italic> and LE<italic>tef1</italic> gene trees
(Figs <xref rid="fig2" ref-type="fig">2</xref>,
<xref rid="fig3" ref-type="fig">3</xref>). This point of discordance
between the gene trees establishes a phylogenetic species limit for isolates
of <italic>H. sulphurea</italic>. In all three gene trees, isolates of <italic>H.
victoriensis</italic> from Australia are phylogenetically distinct from isolates
of <italic>H. sulphurea</italic>. The phylogenetic position of <italic>Protocrea
farinosa</italic> varies between the gene trees. In ITS
(<xref rid="fig1" ref-type="fig">Fig. 1</xref>) and LE<italic>tef1</italic>(<xref rid="fig3" ref-type="fig">Fig. 3</xref>) gene trees, <italic>P.
farinosa</italic> is basal to other species in <italic>Trichoderma</italic> sect.
<italic>Hypocreanum</italic>. In the <italic>rpb2</italic> gene tree, <italic>P. farinosa</italic>resides in the <italic>H. pseudostraminea</italic> clade
(<xref rid="fig2" ref-type="fig">Fig. 2</xref>) with no bootstrap
support. Consequently, the exact phylogenetic position <italic>P. farinosa</italic> in
relation to <italic>Hypocrea</italic> spp. with anamorphs referable to sect.
<italic>Hypocreanum</italic>, is unresolved. Nevertheless, <italic>P. farinosa</italic> is
clearly situated in <italic>Hypocrea s.s</italic>.
(<xref rid="fig5" ref-type="fig">Fig. 5</xref>, clades A2+B2), with a
bootstrap score of 100 uniting the clades, and it will be referred to as
<italic>Hypocrea farinosa</italic> in the remaining sections of this text (see
Taxonomy).</p><p>All three datasets have similar homoplasy indices. The heuristic search of
the most parsimonious tree for the ITS dataset yielded 3193 trees with 217
steps. The minimal possible tree length is 169; the homoplasy index (HI) is
0.221 (<xref rid="fig1" ref-type="fig">Fig. 1</xref>). From 550 total
characters, 421 characters are constant: 41 variable characters are
parsimony-uninformative and 88 characters are parsimony-informative. The
heuristic search of the most parsimonious trees for the <italic>rpb2</italic> dataset
resulted in a tree with 650 steps with the minimum possible tree length of
408: HI = 0.372 (<xref rid="fig2" ref-type="fig">Fig. 2</xref>). From
956 total characters, 651 characters are constant: 88 variable characters are
parsimony-uninformative, and 217 characters are parsimony-informative. The
heuristic search of the most parsimonious trees for the LE<italic>tef1</italic> data
set yielded 64 trees with 314 steps with the minimum possible tree length of
178: HI = 0.433 (<xref rid="fig3" ref-type="fig">Fig. 3</xref>). From
863 total characters, 705 characters are constant, 35 variable characters are
parsimony-uninformative, and 123 characters are parsimony-informative.</p><p>The combined phylogenetic analysis using ITS, partial sequences of
LE<italic>tef1</italic> and <italic>rpb2</italic> showed that <italic>H. sulphurea, H.
subsulphurea, H. victoriensis, H. farinosa</italic>, and <italic>H. corticioides</italic>represent phylogenetically distinct species. <italic>Hypocrea sulphurea, H.
victoriensis, H. subsulphurea</italic>, and <italic>H. corticioides</italic> formed a
monophyletic clade C, supported by a bootstrap score of 77 %, with <italic>H.
sulphurea</italic> distinguished from <italic>H. victoriensis</italic> by a bootstrap
score of 100 % (<xref rid="fig4" ref-type="fig">Fig. 4</xref>).
European and North American isolates of <italic>H. sulphurea</italic> formed a
distinct subclade supported by bootstrap scores of 92 % in the combined
analysis (<xref rid="fig4" ref-type="fig">Fig. 4</xref>), but more
European isolates must be sequenced before determining whether European
isolates represent a distinct phylogenetic species. <italic>Hypocrea citrina, H.
americana, H. pulvinata</italic>, and <italic>H. protopulvinata</italic> formed a strongly
supported monophyletic clade B with a bootstrap score of 100 %
(<xref rid="fig4" ref-type="fig">Fig. 4</xref>). <italic>Hypocrea
microcitrina</italic> and <italic>H. pseudostraminea</italic> are located in an unresolved
clade A, sister to <italic>H. citrina</italic>, supported by a bootstrap score of 90
%. The heuristic search of the most parsimonious trees yielded three trees
with 1202 steps, with the minimum possible tree length of 753: HI = 0.374
(<xref rid="fig4" ref-type="fig">Fig. 4</xref>). From 2358 total
characters, 1767 characters are constant: 163 variable characters are
parsimony-uninformative, and 428 characters are parsimony-informative.</p><p><fig position="float" id="fig2"><label>Fig. 2.</label><caption><p>Parsimony analysis of partial sequences of <italic>rpb2</italic>. Single most
parsimonious tree; 650 steps; consistency index 0.628; retention index 0.771;
homoplasy index 0.372, rest as <xref rid="fig1" ref-type="fig">Fig.
1</xref>.</p></caption><graphic xlink:href="39fig2"></graphic></fig></p><p>The LE<italic>tef1</italic> and <italic>rpb2</italic> regions distinguished between North
American and European isolates of <italic>H. sulphurea</italic>, whereas ITS did not.
The LE<italic>tef1</italic> gene region was less variable than <italic>tef1</italic> sequences
generated by Overton <italic>et al</italic>.
(<xref ref-type="bibr" rid="ref12">2006</xref>), using the primer pair
ef-1/2, for <italic>H. citrina</italic> and allies. Sequences were generated from the
<italic>tef1</italic> gene region for selected species of <italic>H. sulphurea</italic> and
allies included in this study and deposited in GenBank
(<xref ref-type="table" rid="tbl3">Table 3</xref>). The introns of
<italic>tef1</italic> were highly variable making alignments between species such as
<italic>H. citrina</italic> and <italic>H. sulphurea</italic> problematic. Consequently, the
<italic>tef1</italic> region was excluded from this study.</p><p><table-wrap position="float" id="tbl3"><label>Table 3.</label><caption><p>Additional <italic>tef1</italic> sequences deposited in GenBank.</p></caption><table frame="hsides" rules="groups"><thead><tr><th colspan="1" rowspan="1" align="left" valign="top"><bold>Name</bold></th><th colspan="1" rowspan="1" align="left" valign="top"><bold>Strain Number</bold></th><th colspan="1" rowspan="1" align="left" valign="top"><bold>Origin</bold></th><th colspan="1" rowspan="1" align="left" valign="top"><bold>GenBank accession number (<italic>tef1</italic>, primers ef-1, ef-2)</bold></th></tr></thead><tbody><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>H. sulphurea</italic> (Schw.) Sacc
</td><td colspan="1" rowspan="1" align="left" valign="top"> G.J.S. 95-190
</td><td colspan="1" rowspan="1" align="left" valign="top"> Indiana, U.S.A.
</td><td colspan="1" rowspan="1" align="left" valign="top"> DQ835448
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"></td><td colspan="1" rowspan="1" align="left" valign="top"> G.J.S. 95-140
</td><td colspan="1" rowspan="1" align="left" valign="top"> Styria, Austria
</td><td colspan="1" rowspan="1" align="left" valign="top"> DQ835499
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"></td><td colspan="1" rowspan="1" align="left" valign="top"> G.J.S. 00-172
</td><td colspan="1" rowspan="1" align="left" valign="top"> Moscow, Russia
</td><td colspan="1" rowspan="1" align="left" valign="top"> DQ835495
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"></td><td colspan="1" rowspan="1" align="left" valign="top"> G.J.S. 95-176
</td><td colspan="1" rowspan="1" align="left" valign="top"> Kentucky, U.S.A.
</td><td colspan="1" rowspan="1" align="left" valign="top"> DQ835498
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"></td><td colspan="1" rowspan="1" align="left" valign="top"> B.E.O. 98-44
</td><td colspan="1" rowspan="1" align="left" valign="top"> Pennsylvania, U.S.A.
</td><td colspan="1" rowspan="1" align="left" valign="top"> DQ835496
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"></td><td colspan="1" rowspan="1" align="left" valign="top"> B.E.O. 98-45
</td><td colspan="1" rowspan="1" align="left" valign="top"> Pennsylvania, U.S.A.
</td><td colspan="1" rowspan="1" align="left" valign="top"> DQ835497
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>H. victoriensis</italic> Overton
</td><td colspan="1" rowspan="1" align="left" valign="top"> G.J.S. 99-200
</td><td colspan="1" rowspan="1" align="left" valign="top"> Victoria, Australia
</td><td colspan="1" rowspan="1" align="left" valign="top"> DQ835500
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"></td><td colspan="1" rowspan="1" align="left" valign="top"> G.J.S. 99-201
</td><td colspan="1" rowspan="1" align="left" valign="top"> Victoria, Australia
</td><td colspan="1" rowspan="1" align="left" valign="top"> DQ835501
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>H. eucorticioides</italic> Overton
</td><td colspan="1" rowspan="1" align="left" valign="top"> G.J.S. 99-61
</td><td colspan="1" rowspan="1" align="left" valign="top"> Limon, Costa Rica
</td><td colspan="1" rowspan="1" align="left" valign="top"> DQ835502
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>H. farinosa</italic> Berk. & Broome
</td><td colspan="1" rowspan="1" align="left" valign="top">G.J.S. 91-101
</td><td colspan="1" rowspan="1" align="left" valign="top">Maryland, U.S.A.
</td><td colspan="1" rowspan="1" align="left" valign="top">DQ835503
</td></tr></tbody></table></table-wrap></p><p>Species of <italic>Hypocrea</italic> with anamorphs assignable to
<italic>Trichoderma</italic> sect. <italic>Hypocreanum</italic> did not form a monophyletic
group. The K-H test on the combined LE<italic>tef1</italic> and <italic>rpb2</italic> dataset
indicated a significantly worse tree (p < 0.0001) when all
<italic>Hypocrea</italic> with anamorphs in <italic>Trichoderma</italic> sect.
<italic>Hypocreanum</italic> were constrained to monophyly (Monophyletic
<italic>Hypocreanum</italic>, <xref ref-type="table" rid="tbl2">Table 2</xref>).
When taxa with hyphal stromata were constrained to monophyly (Monophyletic
Hyphal, <xref ref-type="table" rid="tbl2">Table 2</xref>) the –log
likelihood was significantly worse (P < 0.0001) than that of the
unconstrained tree.</p><p>The phylogenetic relationship of <italic>Trichoderma</italic> sect. <italic>Pachybasium
s.l.</italic> to clades A2, B2, and C2 could not be established. <italic>Hypocrea
megalocitrina</italic> is situated in clade A2, which was supported by a bootstrap
score of 93 %. <italic>Hypocrea avellanea</italic> and <italic>H. pezizoides</italic>, both of
which have a verticillium-like anamorph, reside in the <italic>H. rufa</italic> clade
C2 (<xref rid="fig5" ref-type="fig">Fig. 5</xref>) supported by a
bootstrap score of 75 %. <italic>Hypocrea parmastoi</italic> and <italic>H.
alcalifuscescens</italic> are located in the unresolved clades F2 and G2, basal to
all species of <italic>Hypocrea</italic> included in this analysis
(<xref rid="fig5" ref-type="fig">Fig. 5</xref>), but have
verticillium-like anamorphs referable to <italic>Trichoderma</italic> sect.
<italic>Hypocreanum</italic>. Based on this dataset, it is unclear whether
<italic>Hypocrea cinereoflava, H. parmastoi</italic>, and <italic>H. alcalifuscescens</italic>should be maintained within <italic>Hypocrea</italic>, as all three species were basal
to other members of the genus (<xref rid="fig5" ref-type="fig">Fig.
5</xref>). For the combined LE<italic>tef1</italic> and <italic>rpb2</italic> dataset,
the heuristic search of the most parsimonious trees yielded three trees with
2469 steps with the minimal possible tree length of 875: HI = 0.646
(<xref rid="fig5" ref-type="fig">Fig. 5</xref>). From 1588 total
characters, 1009 characters were constant, 127 variable characters were
parsimony-uninformative, and 452 characters were parsimony-informative. ML
analysis of the combined data resulted in two trees with log Likelihood scores
of –12767.00537 (not shown). These trees did not significantly differ
from the tree generated in the MP analysis
(<xref rid="fig5" ref-type="fig">Fig. 5</xref>).</p></sec></sec><sec><title>DISCUSSION</title><sec><title>Species recognition</title><p>The combined phylogenetic analyses using ITS and partial sequences of
LE<italic>tef1</italic> and <italic>rpb2</italic> show that <italic>Hypocrea sulphurea, H.
subsulphurea, H. victoriensis, H. farinosa</italic>, and <italic>H.
eucorticioides</italic> represent phylogenetically distinct species that are
members of a strongly supported clade C (Figs
<xref rid="fig4" ref-type="fig">4</xref>,
<xref rid="fig5" ref-type="fig">5</xref>). Dingley
(<xref ref-type="bibr" rid="ref2">1956</xref>) suggested that
morphology could not be used to distinguish between <italic>H. subcitrina</italic> and
<italic>H. sulphurea</italic> and considered these species synonymous. Based on type
studies, we found the ascospores of <italic>H. subcitrina</italic> to be consistently
shorter and narrower than those of <italic>H. sulphurea.</italic> In contrast,
<italic>Hypocrea egmontensis</italic> is considered a facultative synonym of the older
<italic>H. subcitrina.</italic></p><p>Dingley deposited a culture of <italic>H. sulphurea</italic>(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=500.67&link_type=CBS">CBS 500.67</ext-link>) from
New Zealand in CBS. Specimens recently collected from Australia had the same
ITS sequence as <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=500.67&link_type=CBS">CBS
500.67</ext-link> and represent Dingley's concept of <italic>H. sulphurea</italic>.
Molecular phylogenetic results indicate that the Australian specimens and the
New Zealand culture (<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=500.67&link_type=CBS">CBS
500.67</ext-link>) represent a new phylogenetic species, described here as
<italic>H. victoriensis</italic>, that differs morphologically from <italic>H.
subcitrina</italic> and <italic>H. sulphurea</italic>. The morphological similarities
between Australian specimens of <italic>H. victoriensis</italic> and North American
specimens of <italic>H. sulphurea</italic> are striking, but the part-ascospores of
the Australian species are more strongly spinulose than the part-ascospores
found in <italic>H. sulphurea</italic>. In addition, none of the Australian specimens
occurred on <italic>Exidia</italic> spp., which is a common substrate in North
America. This suggests that ascospore ornamentation and substratum are
informative species characters for members of the <italic>H. sulphurea</italic>subclade (clade C, <xref rid="fig4" ref-type="fig">Fig. 4</xref>).</p></sec><sec><title>Hyphal versus pseudoparenchymatous stromata</title><p><italic>Hypocrea</italic> species with hyphal stromata and anamorphs assignable to
<italic>Trichoderma</italic> sect. <italic>Hypocreanum</italic> are situated in different
clades. <italic>Hypocrea megalocitrina</italic> resides in clade A2
(<xref rid="fig5" ref-type="fig">Fig. 5</xref>) with <italic>H.
psychrophila. Hypocrea avellanea</italic> has a hyphal stroma and a
verticillium-like anamorph with conidia that are uniform in size and shape.
Anamorphs in <italic>Trichoderma</italic> sect. <italic>Hypocreanum</italic> typically produce
conidia that are variable in size and shape. <italic>Hypocrea avellanea</italic>resides in the <italic>H. rufa</italic> clade (<xref rid="fig5" ref-type="fig">Fig.
5</xref>) with species having pseudoparenchymatous stromata. Anamorphs in
the <italic>Hypocrea rufa</italic> clade generally produce conidia that are typically
more uniform in size and shape than those found in <italic>Trichoderma</italic> sect.
<italic>Hypocreanum. Hypocrea alcalifuscescens</italic> and <italic>H. parmastoi</italic> have
hyphal stromata and verticillium-like anamorphs and are basal to other major
clades of <italic>Hypocrea/Trichoderma</italic>. Species found in clade B2
(<xref rid="fig5" ref-type="fig">Fig. 5</xref>), have effused,
extensive stromata, with pseudoparenchymatous tissue, except one, <italic>H.
subsulphurea,</italic> which is hyphal. Anamorphs in clade B2 produce conidia
variable in size and shape, typical of <italic>Trichoderma</italic> sect.
<italic>Hypocreanum. Hypocrea pezizoides</italic>, known to have a
pseudoparenchymatous stroma and a verticillium-like anamorph also resides in
the <italic>H. rufa</italic> clade (C2, <xref rid="fig5" ref-type="fig">Fig.
5</xref>), a finding consistent with Chaverri <italic>et al.</italic>(<xref ref-type="bibr" rid="ref1">2003</xref>). The anamorph of <italic>H.
pezizoides</italic> produces conidia that initially are light green, but become
hyaline after repeated transfers. Species with pseudoparenchymatous stroma and
anamorphs that produce hyaline conidia variable in size and shape are located
in clade B2 (<xref rid="fig5" ref-type="fig">Fig. 5</xref>). Species
with hyphal stromata and anamorphs that produce uniform conidia (of similar
size and shape) are polyphyletic.</p><p><fig position="float" id="fig3"><label>Fig. 3.</label><caption><p>Parsimony analysis of partial sequences of LE<italic>tef1</italic>. One phylogram
of 64 most parsimonious trees; 314 steps; consistency index: 0.567; retention
index: 0.744; homoplasy index: 0.433, rest as
<xref rid="fig1" ref-type="fig">Fig. 1</xref>.</p></caption><graphic xlink:href="39fig3"></graphic></fig></p><p><fig position="float" id="fig4"><label>Fig. 4.</label><caption><p>Combined parsimony analysis of ITS, LE<italic>tef1</italic>, <italic>rpb2</italic>.
Phylogram of one of three most parsimonious trees; 428 steps; consistency
index: 0.626; retention index: 0.766; homoplasy index: 0.374, rest as
<xref rid="fig1" ref-type="fig">Fig. 1</xref>.</p></caption><graphic xlink:href="39fig4"></graphic></fig></p><p>Petch (<xref ref-type="bibr" rid="ref13">1937</xref>) established
the genus <italic>Protocrea</italic> Petch for species that have simple ascomata
immersed or seated upon a byssoid stroma with ascospores that disarticulate
into part-ascospores. Rossman <italic>et al.</italic>(<xref ref-type="bibr" rid="ref15">1999</xref>) described the anamorph
of <italic>Protocrea</italic> as acremonium- or verticillium-like. <italic>Protocrea
farinosa</italic> resides in clade B2 (<xref rid="fig5" ref-type="fig">Fig.
5</xref>) with other species with acremonium- and verticillium-like
anamorphs. A well-defined layer of pseudoparenchymatous tissue was observed
below the perithecia in specimens of <italic>P. farinosa</italic>. Although the
teleomorphs of specimens examined varied in the degree of pseudoparenchymatous
tissue present, the part-ascospore measurements obtained are identical to
those published for <italic>P. farinosa</italic> by Rossman <italic>et al</italic>.
(<xref ref-type="bibr" rid="ref15">1999</xref>) and the anamorph
characteristics are identical to those described by Doi
(<xref ref-type="bibr" rid="ref5">1972</xref>) for <italic>P.
farinosa</italic>.</p></sec><sec><title>Trichoderma sect. <italic>Hypocreanum</italic> and classification</title><p>The phylogeny of the major clades in <italic>Trichoderma</italic>/<italic>Hypocrea</italic>is essentially unresolved based on the genes used in this study. However,
<italic>Hypocrea</italic> spp. with well-defined pseudoparenchymatous stroma tissue,
and acremonium- or verticillium-like conidiophores (hypocreanum-like), that
produce hyaline conidia variable in size and shape, can be accommodated in a
large monophyletic assemblage of species B2
(<xref rid="fig5" ref-type="fig">Fig. 5</xref>). Kullnig-Gradinger
<italic>et al.</italic> (<xref ref-type="bibr" rid="ref11">2002</xref>)
suggested that <italic>Trichoderma</italic> sect. <italic>Hypocreanum</italic> and sect.
<italic>Pachybasium</italic> should be merged as they are phylogenetically
indistinguishable. The present study, which included 17 taxa morphologically
belonging to sect. <italic>Hypocreanum</italic>, shows that the phylogenetic
relationship of <italic>Trichoderma</italic> sect. <italic>Hypocreanum</italic> to sect.
<italic>Pachybasium</italic> could not be resolved in the combined LE<italic>tef1</italic> and
<italic>rpb2</italic> dataset. The anamorphs of <italic>H. megalocitrina, H.
parmastoi</italic>, and <italic>H. alcalifuscescens</italic> are morphologically similar
to anamorphs typical of <italic>Trichoderma</italic> sect. <italic>Hypocreanum</italic>;
nevertheless, these fungi do not belong to the major <italic>Hypocreanum</italic>clade B2 (<xref rid="fig5" ref-type="fig">Fig. 5</xref>), nor are they
phylogenetically related to members of <italic>Trichoderma</italic> sect.
<italic>Pachybasium</italic>.</p><p><fig position="float" id="fig5"><label>Fig. 5.</label><caption><p>Parsimony analysis of the combined LE<italic>tef1</italic> and <italic>rpb2</italic> data
set. Phylogram of one of 3 most parsimonious trees; 2469 steps; consistency
index: 0.354; retention index: 0.515; homoplasy index: 0.646, rest as
<xref rid="fig1" ref-type="fig">Fig. 1</xref>. Outgroup taxa:
<italic>Hypomyces stephanomatis</italic>; <italic>Acrachnocrea scabrida</italic>;<italic> Sphaerostibella</italic> cf. <italic>aureonitens</italic>.</p></caption><graphic xlink:href="39fig5"></graphic></fig></p><p>The multigene phylogeny of Kullnig-Gradinger <italic>et al.</italic>(<xref ref-type="bibr" rid="ref11">2002</xref>) should serve as an
example for future phylogenetic analyses to determine sectional relationships,
but future studies should include a larger number of taxa and exclude ITS rDNA
sequences. The ITS region proved useful in distinguishing between closely
related species (<xref ref-type="bibr" rid="ref12">Overton <italic>et al.</italic>2006</xref>) and has been used for the revision of sections
<italic>Longibrachiatum</italic> and <italic>Trichoderma</italic> (Kuhls <italic>et al.</italic><xref ref-type="bibr" rid="ref9">1996</xref>,
<xref ref-type="bibr" rid="ref10">1997</xref>; Kinderman <italic>et
al.</italic> 1998; Samuels <italic>et al.</italic><xref ref-type="bibr" rid="ref18">1998</xref>,
<xref ref-type="bibr" rid="ref17">1999</xref>).</p><p>Overton <italic>et al.</italic>(<xref ref-type="bibr" rid="ref12">2006</xref>) demonstrated that ITS
rDNA, <italic>rpb2</italic>, and the <italic>tef1</italic> region could establish phylogenetic
species limits, but the introns found in the <italic>tef1</italic> region, delimited
by the primers ef-1 and ef-2, were highly divergent among morphologically
similar species. In this study partial sequences of the large exon
(LE<italic>tef1</italic>) were generated for the seven species, including several of
those treated by Overton <italic>et al.</italic>(<xref ref-type="bibr" rid="ref12">2006</xref>). The LE<italic>tef1</italic>region also resolved all major clades established by these authors using the
<italic>tef1</italic> gene region and distinguished between North American and
European isolates of <italic>H. pulvinata</italic>; therefore LE<italic>tef1</italic> is
better suited for phylogenetic studies than the <italic>tef1</italic> region
previously sequenced by Overton <italic>et al.</italic>(<xref ref-type="bibr" rid="ref12">2006</xref>).</p><p>Comparatively few of the approximately 200 named species of
<italic>Hypocrea</italic> have been sequenced to date, with published accounts placing
an over-reliance on ITS rDNA sequence data. The LE<italic>tef1</italic> and
<italic>rpb2</italic> sequences generated in this study, work by Chaverri <italic>et
al.</italic> (<xref ref-type="bibr" rid="ref1">2003</xref>), and data from
other gene regions published by Kullnig-Gradinger <italic>et al.</italic>(<xref ref-type="bibr" rid="ref11">2002</xref>), have helped to clarify
our understanding of the sectional relationships of
<italic>Hypocrea/Trichoderma</italic>. Additional taxa from other genera such as
<italic>Sarawakus</italic> Lloyd, with <italic>Trichoderma</italic> anamorphs, need to be
sequenced before a complete phylogeny of <italic>Hypocrea/Trichoderma</italic> can be
established.</p></sec><sec><title>The evolution of anamorphs referable to <italic>Trichoderma</italic> sect.
<italic>Hypocreanum</italic></title><p>There has been considerable speculation published on the evolution of the
anamorphs in <italic>Trichoderma</italic> sect. <italic>Hypocreanum.</italic> Samuels
(<xref ref-type="bibr" rid="ref16">1996</xref>) hypothesized that the
anamorphs referable to <italic>Trichoderma</italic> sect. <italic>Hypocreanum</italic> may be
synanamorphs or spermatial states, suggesting that <italic>Hypocrea</italic> with
acremonium- or verticillium-like anamorphs with hyaline conidia have lost the
ability to produce a primary trichoderma-like anamorph, with pyramidally
branched conidiophores and green conidia. Kullnig-Gradinger <italic>et al.</italic>(<xref ref-type="bibr" rid="ref11">2002</xref>) presented molecular
data suggesting that the more typical trichoderma-like anamorph with green
conidia may have evolved from genera having verticillium-like anamorphs, in
particular <italic>Aphysiostroma</italic> Barrasa and <italic>Arachnocrea</italic> Z.
Moravec.</p><p>Results based on molecular data have not conclusively established the
evolution of the <italic>Trichoderma</italic> anamorph, including those referable to
sect. <italic>Hypocreanum</italic>. Two species are of particular interest when
considering the hypotheses promulgated by Kullnig-Gradinger <italic>et al.</italic>(<xref ref-type="bibr" rid="ref11">2002</xref>) and Samuels
(<xref ref-type="bibr" rid="ref16">1996</xref>). <italic>Hypocrea
pezizoides</italic> has light green conidia that become completely hyaline in
subsequent transfers, suggesting an incomplete reversal to the primitive
verticillium-like form with hyaline conidia. This species resides in the
<italic>H. rufa</italic> clade C2 (<xref rid="fig5" ref-type="fig">Fig.
5</xref>) based on combined <italic>rpb2</italic> and LE<italic>tef1</italic> gene
sequences and based on ITS sequence data, a finding consistent with
Kullnig-Gradinger <italic>et al</italic>.
(<xref ref-type="bibr" rid="ref11">2002</xref>). <italic>Hypocrea
cinereoflava</italic> produces a primary synnematous anamorph and a
verticillium-like synanamorph. This species of <italic>Hypocrea</italic> is important
when considering the hypothesis of Samuels
(<xref ref-type="bibr" rid="ref16">1996</xref>) that the
verticillium-like anamorphs found in <italic>Trichoderma</italic> sect.
<italic>Hypocreanum</italic> represent spermatial states, in which the primary
trichoderma-like anamorph was lost. <italic>Hypocrea cinereoflava</italic> is located
in an unresolved basal clade of <italic>Hypocrea s.s.</italic>(<xref rid="fig5" ref-type="fig">Fig. 5</xref>) and, based on the
molecular results of this study, it could not be excluded from the genus
<italic>Hypocrea</italic>. The phylogenetic placement of this species basal to all
other <italic>Hypocrea</italic> species sequenced in this analysis could suggest that
the ability to produce a synnematous primary anamorph has subsequently been
lost. The data obtained in this study provide some support for the hypotheses
of Samuels (<xref ref-type="bibr" rid="ref16">1996</xref>) and
Kullnig-Gradinger <italic>et al</italic>.
(<xref ref-type="bibr" rid="ref11">2002</xref>), leaving room for
speculation. Additional taxa need to be sequenced before the evolution of
<italic>Trichoderma</italic> anamorphs can be more accurately determined.</p></sec></sec><sec><title>TAXONOMY</title><p><list list-type="order"><list-item><p><italic><bold>Hypocrea sulphurea</bold></italic> (Schw.) Sacc., Syll. Fung. 2: 535. 1883.
Figs <xref rid="fig6" ref-type="fig">6</xref>,
<xref rid="fig7" ref-type="fig">7</xref>,
<xref rid="fig8" ref-type="fig">8</xref>.</p><p><list list-type="simple"><list-item><p>≡ <italic>Sphaeria sulphurea</italic> Schw., Trans. Amer. Philos. Soc. 2:
193. 1832.</p></list-item><list-item><p>= <italic>Hypocrea sulphurea</italic> f. <italic>macrospora</italic> Yoshim. Doi, Bull.
Natl. Sci. Mus. 15: 699. 1972.</p></list-item></list></p><p><italic>Anamorph</italic>: <italic>Trichoderma</italic> sp. [sect.
<italic>Hypocreanum</italic>].</p><p><italic>Teleomorph</italic>: Stromata effuse, extensive, largest continuous stroma
70 × 30 mm, smallest continuous stroma 1 × 1 mm, many stromata not
larger than 25 × 10 mm, varying in colour, sometimes vivid yellow,
usually light yellow to greyish yellow (3A8; 3A5–3A6; 4A5–4A6),
KOH<sup>+/–</sup>, reaction variable, usually very weak, the stroma
becoming light orange (6A4); ostiolar canals visible at the stroma surface,
appearing light orange (6A4), giving rise to the greyish yellow overall
appearance of the stroma. Stroma surface smooth; tissue immediately below the
stroma surface formed of compact to loose pseudoparenchymatous cells of
<italic>textura globulosa</italic> to <italic>t. angularis</italic>. Perithecia completely
immersed, generally widely spaced, compact in some regions, sometimes
completely absent near the margins or regions of extensive stroma growth.
Perithecia ellipsoidal, (128–)190–250(–277) μm long
(including the length of the ostiolar canal, n = 26); width of perithecia near
the base (measured from 3/4 total length of the perithecium),
(87–)100–142(–175) μm (n = 26); length of ostiolar canal
(42–)52–74(–85) μm; width of ostiolar canal from outer
perithecial wall to the opposite internal perithecial wall
(22–)30–50(–64) μm (n = 26); wall
KOH<sup>+/–</sup>, reaction variable, weak. Asci cylindrical,
(80–)94–116(–150) ×
(4.2–)5.3–7.1(–8.3) μm (n = 196); tip slightly thickened.
Part-ascospores hyaline, thick-walled, spinulose, dimorphic; distal part
obovate, sometimes subellipsoidal, (4.2–)5.2–6.6 (–7.6)
× (4.2–)5.3–7.1(–8.3) μm, L/W ratio,
(0.8–)1.1–1.4 (–1.7) (n = 294); proximal part ellipsoidal,
sometimes subcylindrical, (4.4–)5.5–6.9(–8.5) ×
(2.7–)3.9–5.1 (–6.6) μm, L/W ratio
(0.9–)1.2–1.6(–2.2) (n = 294).</p><p><italic>Anamorph</italic>: Colonies covering a 100 mm diam Petri plate with PDA in
10 d, not producing concentric rings or radial rays of mycelium; a layer of
aerial mycelium covering the entire Petri plate, aerial mycelium consisting of
visible conidiophores; conidiophores irregularly branched, on long hyphal
elements, usually verticillium-like; phialides in whorls of 2–4,
solitary or alternating in pairs on long hyphal elements; phialides subulate,
(8–)17–32(–45) × (2.4–)3–4(–4.8)
μm (n = 92); conidia variable in size, obovate to subcylindrical, often
ellipsoidal, (3.9–) 5.6–9.0(–12.6) ×
(3.0–)3.3–4.3(–6.6) μm (n = 112), with some conidia
asymmetric, having a flat edge; no distinctive odour; yellowish orange pigment
(4A6–4A8) produced near the inoculation point. After 10 d conidia
beginning to swell and more variable in size. Colonies on SNA or CMD did not
produce conidiophores in 10 d.</p><p><italic>Habitat</italic>: Found on decorticated wood with <italic>Exidia</italic> spp.,
sometimes occurring on decorticated wood without visible evidence of
<italic>Exidia</italic> spp.</p><p><italic>Known distribution</italic>: Europe, Japan and North America.</p><p><italic>Isotype</italic>: <bold>U.S.A.,</bold> Pennsylvania, Salem & Bethlehem, on
<italic>Exidia</italic> sp., <italic>H. sulphurea</italic> (K, herb. Schweinitz).</p><p><italic>Other specimens examined</italic>: <bold>Austria,</bold> Styria, Leibnitz, St.
Nikolai, alt. 310 m, on decorticated wood, <italic>Exidia</italic> sp. not visible, 26
Aug. 1995, H. Voglmayr (BPI 737705; culture G.J.S. 95-140). <bold>Japan,</bold>Amori Prefecture, near Tsuta-Onsen, Towanda National Park, Towanda-Cho,
Kami-kita-Gun, on <italic>Exidia</italic> sp., 10 Sep. 1971, Y. Doi, (NY, TNS. D-1169
= TNS-F-190169), <bold>paratype</bold> of <italic>H. sulphurea</italic> f.
<italic>macrospora</italic>. <bold>Russia,</bold> 10 km northeast of Moscow, mixed deciduous
forest, 17 Oct. 2000, A. Alexandrova (BPI 748252; culture G.J.S. 00-172).
<bold>U.S.A.,</bold> Indiana, Brown County, vic. Pikes Peak, Happy Hollow Camp, alt.
250 m, 39°09' N, 86°06' W, on bark with unidentified fungus, 29 Sep.
1995, G. J. Samuels (BPI 737764; culture G.J.S. 95-190); Brown County, Yellow
Wood State Forest, Jackson Creek Management Trail, alt. 200 m, 39°09' N,
86°06' W, on <italic>Exidia</italic> sp., 30 Sep. 1995, G.J. Samuels (BPI 737772;
culture G.J.S. 95-198); Illinois, Carbondale, Giant City State Park, on
<italic>Exidia</italic> sp., 9 Aug. 1999, B.E. Overton, B.E.O. 99-02 (BPI); Union
County, Carbondale, Giant City State Park, on leaf litter and decorticated
wood, 19 Sep. 1994, G. J. Samuels (BPI 749353; culture G.J.S. 94-58);
Kentucky, Rowan County, Daniel Boone National Forest, Cave Run Lake, Sheltowee
Trail, on <italic>Exidia</italic> sp., 26 Sep. 1995, G. J. Samuels (BPI 737752;
culture G.J.S. 95-176); Maryland, Prince Georges County, Laurel, Patuxent
Refuge, on <italic>Exidia</italic> sp., 2 July 2000, Kadri Põldmaa, K.P. 00-14
(BPI; TFC 2000-52); Tacoma Park, on <italic>Exidia</italic> sp., Dec. 1906, C.L. Shear
(BPI 631489); New York, Green County, on bark, no <italic>Exidia</italic> sp. visible,
27 Sep. 1998, B.E. Overton, B.E.O. 98-50 (BPI); North Carolina, Durham County,
Hill Forest, on <italic>Carya glabra</italic> var. <italic>glabra</italic> with
<italic>Exidia</italic> sp., 18 May 2002, L. Grand (NCSU Mycological Herbarium); North
Dakota, Fargo, on branches of <italic>Tilia americana</italic> with <italic>Exidia</italic>sp., 1907–1908, G. W. Wilson & F. J. Seaver (BPI 631488);
Pennsylvania, Center County, Rock Springs Agricultural Research Center, on
<italic>Exidia</italic> sp., 19 Sep. 1998, B.E. Overton, B.E.O. 98-44 (BPI); same
origin B.E.O. 98-45 (BPI); Vermont, Burlington, Indian Brook Conservation
Area, Aug. 2000, B.E. Overton, B.E.O. 00-07 (BPI; culture G.J.S. 00-76).</p><p><italic>Comments</italic>: The paratype specimen of <italic>H. sulphurea</italic> f.
<italic>macrospora</italic> and the specimens from Russia and Austria had
part-ascospores that were on average 1 μm larger than specimens of <italic>H.
sulphurea</italic> from North America; <italic>H. sulphurea</italic> f.
<italic>macrospora</italic> and European specimens (Russia and Austria) had distal
part-ascospores, (5.6–) 6.0–7.1(–7.6) ×
(4–)4.8–5.9(–6.5) μm, and proximal part-ascospores,
(5.6–)6.4–7.6(–8.5) ×
(3.5–)4.3–5.7(–6.6) μm; <italic>H. sulphurea</italic> specimens
from North America had distal part-ascospores
(4.2–)5.3–6.4(–7.1) × (3.6–)
4.2–5(–5.8) μm, and proximal part-ascospores (4.4–)
5.3–6.4(–8.2) × (2.7–)3.9–4.7(–5.7)
μm.</p><p>European and North American isolates were slightly different in
LE<italic>tef1</italic> and <italic>rpb2</italic> gene trees, but in the ITS tree one European
isolate grouped with isolates of <italic>H. sulphurea</italic> from North America. We
use this point of discordance to establish the phylogenetic species limit for
<italic>H. sulphurea. Hypocrea sulphurea</italic> f. <italic>macrospora</italic> is not
considered sufficiently distinct from <italic>H. sulphurea</italic>. The teleomorph
description provided above for <italic>H. sulphurea</italic> consists of combined
measurements from all specimens examined for this species. Even with variable
part-ascospores, the ascospores of <italic>H. sulphurea</italic> are significantly
larger than those of <italic>H. subcitrina</italic>, even at the lower extremes;
therefore the synonymy proposed by Dingley 1956 is rejected.</p><p>Doi (<xref ref-type="bibr" rid="ref5">1972</xref>) described an
additional species <italic>H. megalosulphurea</italic> Yoshim. Doi in which proximal
part-ascospores can be as large as 10 μm diam. Type material or cultures
were not available for study, but it is doubtful that <italic>H.
megalosulphurea</italic> is a synonym of <italic>H. sulphurea</italic> because, even
though part-ascospores can vary in size, variation of this magnitude was never
observed in the specimens of <italic>H. sulphurea</italic> examined.</p></list-item><list-item><p><italic><bold>Hypocrea subcitrina</bold></italic> Kalchbr. & Cooke, Grevillea 9: 26.
1880)</p><p><list list-type="simple"><list-item><p>= <italic>H. egmontensis</italic> Dingley, Trans. Roy. Soc. New Zealand 83: 647.
1956.</p></list-item></list></p><p><italic>Anamorph</italic>: Unknown.</p><p><italic>Notes</italic>: G.J. Samuels (pers. comm.) provided measurements of
part-ascospores that allow a comparison of type specimens as follows:
<italic>Hypocrea subcitrina</italic>, distal part-ascospores subglobose to obovate
conical, (4.5–)4.7–5.1(–5.4) ×
(3.6–)3.8–4.2(–4.3) μm; proximal part tending to be
oblong and narrow, (4.2–)4.7–5.6(–6.0) ×
(3.6–)3.5–3.7(–3.8) μm; <italic>H. egmontensis</italic>, distal
part-ascospores subglobose, conical, (4.1–)4.5–5.3(–5.7)
× (3.8–)4–4.6(–5.2) μm; proximal part-ascospores
(4.3–)5.0–5.6(–6.7) ×
(2.7–)3.5–4.3(–4.1) μm. In this respect <italic>H.
subcitrina</italic> and <italic>H. egmontensis</italic> are identical, differing from the
larger ascospores of <italic>H. sulphurea</italic> and <italic>H. victoriensis</italic>. Based
on ascospore measurements, <italic>H. egmontensis</italic> is considered a facultative
synonym of the older name <italic>H. subcitrina</italic>. Doi
(<xref ref-type="bibr" rid="ref4">1971</xref>) already suggested that
<italic>H. egmontensis</italic> and <italic>H. subcitrina</italic> were similar species. Doi
(<xref ref-type="bibr" rid="ref4">1971</xref>) described <italic>H.
subcitrina</italic> var. <italic>dimorphospora</italic> Yoshim. Doi. based on the presence
of part-ascospores of different size classes in specimens collected in New
Guinea. Type material was not available for study and the relationship of this
variety to <italic>H. subcitrina</italic> cannot be formally evaluated.</p><p><italic>Specimens examined</italic>: <bold>South Africa,</bold> Port Natal, <italic>H.
subcitrina</italic>, Wood 184 (K; <bold>isotype</bold>). <bold>New Zealand,</bold> Taranaki,
Mt Egmont, Apr. 1946, J.M. Dingley 6272, <italic>H. egmontensis</italic> (PDD 6272;
<bold>holotype</bold>).</p></list-item><list-item><p><italic><bold>H. victoriensis</bold></italic> Overton, <bold>sp. nov</bold>. MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB501055&link_type=MB">MB501055</ext-link>. Figs
<xref rid="fig9" ref-type="fig">9</xref>,
<xref rid="fig10" ref-type="fig">10</xref>.</p><p><italic>Anamorph</italic>: <italic>Trichoderma</italic> sp. [sect.
<italic>Hypocreanum</italic>].</p><p><italic>Etymology</italic>: Named after the location where it was collected,
Victoria, Australia.</p><p>Stromata effusa, extensa, rubido-lutea vel griseo-lutea,
KOH<sup>+/–</sup>. Ascosporae hyalinae, crassitunicatae, spinulosae,
dimorphicae; pars distalis plus minusve ellipsoidea,
(4.8–)5.6–6.8(–7.4) × (3.9–)
4.5–5.5(–5.9) μm, pars proxima
(4.7–)5.8–7.4(–8.6) ×
(3.6–)4.1–5.1(–6.1) μm. Anamorphosis <italic>Trichoderma</italic>sectionis <italic>Hypocreanum</italic>. Conidia hyalina, obovata vel subellipsoidea,
(4.5–)5.9–9.3(–12.0) ×
(2.8–)3.2–4.2(–5.2) μm.</p><p>Typus: BPI 747361.</p><p><italic>Teleomorph</italic>: Stromata effuse, extensive, largest continuous stroma
30 × 10 mm, smallest continuous stroma 3 × 2 mm, varying in
colour, usually reddish yellow to greyish yellow (4A7–4B7),
KOH<sup>+/–</sup>, reaction variable, usually very weak with stroma
becoming light orange (6A4); ostiolar openings visible at the stroma surface,
appearing light orange (6A4), giving rise to the greyish yellow overall
appearance of the stroma. Stroma surface smooth, tissue immediately below the
stromatal surface formed of compact to loose pseudoparenchymatous cells,
<italic>textura globulosa</italic> to <italic>t. angularis</italic>. Perithecia completely
immersed, with ostiolar canals projecting from the stroma surface,
(35–)36–51(–61) μm (n = 10); perithecia generally widely
spaced, compact in some regions, sometimes completely absent near the margins
or regions of extensive stroma growth. Perithecia ellipsoidal,
(335–)345–405(–435) μm high (including the length of the
ostiolar canal, n = 10); width of perithecia near the base (measured from 3/4
total length of the perithecium), (152–)160–205(–225) μm
(n = 10); length of ostiolar canal (100–)105–140(–155)
μm; width of ostiolar canal, from outer perithecial wall to opposite
internal perithecial wall, (38–)41–53(–54) μm (n =10);
wall KOH<sup>+/–</sup>, reaction variable, weak. Asci cylindrical,
(92–)112–142(–152) × (5.5–)6–7(–9)
μm (n = 49); tip slightly thickened. Part-ascospores hyaline, thick-walled,
distinctly spinulose, dimorphic; distal part subellipsoidal, sometimes
obovate, (4.8–)5.6–6.8 (–7.4) ×
(3.9–)4.5–5.5(–5.9) μm, L/W ratio
(1–)1.1–1.3(–1.6) (n = 72); proximal part, ellipsoidal,
sometimes subellipsoidal, (4.7–)5.8–7.4(–8.6) ×
(3.6–)4.1–5.1 (–6.1) μm, L/W ratio
(1–)1.2–1.7(–2) (n = 72).</p><p><italic>Anamorph</italic>: Colonies covering a 100 mm diam Petri plate with PDA in
10 d, not producing concentric rings or radial rays of mycelium; a layer of
aerial mycelium covering the agar surface; conidiophore and conidium
production limited; conidiophores irregularly branched, on long hyphal
elements, usually verticillium-like; phialides not in whorls, solitary or
alternating in pairs on long hyphal elements, subulate,
(3.8–)18–36(–57) ×
(2.6–)3.1–3.9(–4.4) μm (n = 62); conidia variable in
size, obovate to elongate-obovate or subellipsoidal,
(4.5–)5.9–9.3(–12.0) ×
(2.8–)3.2–4.2(–5.2) μm (n = 40); no distinctive odour;
yellowish orange pigment (4A8) produced near the inoculation point. After 10 d
conidia beginning to swell and more variable in size. Colonies on SNA or CMD
did not produce conidiophores within 10 d.</p><p><italic>Habitat</italic>: Typically found on decorticated wood or bark, species of
<italic>Exidia</italic> not observed, possibly fungicolous.</p><p><italic>Known distribution</italic>: Australia, New Zealand.</p><p><italic>Holotype</italic>: <bold>Australia,</bold> Victoria, Otway National Park, along
Great Ocean Road, Cannan's Track, alt. 350 m, on bark, 27 Aug. 1999, G. J.
Samuels (BPI 747361; culture G.J.S. 99-200).</p><p><italic>Other specimens examined</italic>: <bold>Australia,</bold> same origin as
holotype (BPI 747362; culture G.J.S. 99-201); Otway Ranges, Melba Gully State
Park, Madsen Track along the Johanna River, alt. 350 m, on bark, 27 Aug. 1999,
G. J. Samuels (BPI 747363; culture G.J.S. 99-130). <bold>New Zealand</bold>, Culture
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=500.67&link_type=CBS">CBS 500.67</ext-link>, as
“<italic>Hypocrea sulfurea”</italic>, PDD 6332, specimen not located.</p><p><italic>Comments</italic>: Isolate
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=500.67&link_type=CBS">CBS 500.67</ext-link> had the
same ITS sequence as isolates from Australia included here under the name
<italic>H. victoriensis</italic>. The part-ascospore measurements of <italic>H.
victoriensis</italic> and <italic>H. sulphurea</italic> are substantially larger than
those found in the type of <italic>H. subcitrina</italic>. Molecular phylogenetic data
indicate that <italic>H. sulphurea</italic> and <italic>H. victoriensis</italic> are
phylogenetically close, but distinct species. <italic>H. victoriensis</italic> has
part-ascospores that are distinctly more spinulose than those of <italic>H.
sulphurea</italic>. In addition, the ostioles project conspicuously from the
surface in <italic>H. victoriensis</italic> whilst in <italic>H. sulphurea</italic> the
ostioles protrude only slightly. Furthermore, <italic>H. victoriensis</italic> does
not appear to grow on species of <italic>Exidia</italic>.</p></list-item><list-item><p><italic><bold>Hypocrea eucorticioides</bold></italic> Overton, <bold>nom. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB501056&link_type=MB">MB501056</ext-link>. Figs
<xref rid="fig11" ref-type="fig">11</xref>,
<xref rid="fig12" ref-type="fig">12</xref>.</p><p><list list-type="simple"><list-item><p>≡ <italic>Hypocrea corticioides</italic> Speg., An. Mus. Nac. Hist. Nat.
Buenos Aires 23: 75. 1912 [non Berk. & Broome, J. Linn. Soc. Bot. 14: 111.
1873].</p></list-item></list></p><p><italic>Anamorph</italic>: <italic>Trichoderma</italic> sp. [sect.
<italic>Hypocreanum</italic>].</p><p><italic>Teleomorph</italic>: Stromata effuse, extensive, largest continuous stroma
15 × 10 mm, smallest continuous stroma 7 × 1 mm, varying in
colour, typically greyish yellow (4A5–4B5), sometimes pastel-yellow to
light yellow (2A4–2A5), KOH<sup>+/–</sup>, reaction variable,
usually very weak with stroma becoming orange (6A8); ostiolar canals visible
at stroma surface, appearing light orange (6A4), giving rise to the greyish
yellow overall appearance of the stroma. Stroma surface smooth, tissue
immediately below the stromatal surface formed of compact to loose
pseudoparenchymatous cells, <italic>textura globulosa</italic> to <italic>t.
angularis</italic>. Perithecia completely immersed, generally widely spaced,
compact in some regions, sometimes completely absent near the margins or
regions of extensive stroma growth. Perithecia subglobose to subellipsoidal,
(130–)147–193(–200) μm high (including the length of the
ostiolar canal, n = 14); width of perithecia near the base (measured from 3/4
total length of the perithecium) (79–)89–125(–135) μm (n
= 14); length of ostiolar canal (25–)40–57(–60) μm; width
of ostiolar canal from outer perithecial wall to opposite internal perithecial
wall (23–)37–42(–46) μm (n =14); wall
KOH<sup>+/–</sup>, reaction variable, weak. Asci cylindrical,
(56–)62–72(–80) ×
(3.5–)5.1–6.0(–7.3) μm (n = 60); tip slightly thickened.
Part-ascospores hyaline, thin-walled, spinulose, monomorphic; generally
subglobose, (2.6–)2.8–3.5(–3.9) ×
(2.4–)2.6–3.2(–3.7) μm, L/W ratio
(0.8–)0.9–1.2(–1.3) (n = 60).</p><p><italic>Anamorph</italic>: Colonies covering a 100 mm diam Petri plate with PDA in
10 d, not producing concentric rings or radial rays of mycelium; a layer of
aerial mycelium covering the entire plate; conidiophores irregularly branched,
on long hyphal elements, usually verticillium-like; phialides in whorls of
2–3, solitary or alternating on long hyphal elements; phialides
subulate, (4.1–)8–16(–23.2) ×
(1.5–)2.3–3.1(–5.6) μm (n = 34); conidia variable in
size, typically subglobose, but sometimes subellipsoidal,
(2.4–)3.3–4.2(–10) ×
(2.0–)2.5–3.1(–6) μm (n = 19); no distinctive odour or
pigment. Colonies on SNA or CMD did not produce conidiophores within 10 d.</p><p><italic>Habitat</italic>: Typically found on bark of decaying wood.</p><p><italic>Known distribution</italic>: Central and South America.</p><p><italic>Holotype</italic>: <bold>Argentina</bold>, Entre Rios, Ibicuy, 28 June 1911, C.
Spegazzini no. 911 (LPS 1719).</p><p><italic>Other specimens examined</italic>: <bold>Costa Rica,</bold> Limón, Puerto
Viejo, Refugio Nacional Mendaca-Manzanilla, on decorticated wood, July 1999,
G.J. Samuels & P. Chaverri (BPI 747358; culture G.J.S. 99-61).
<bold>Venezuela,</bold> Bolívar, La Urbana, Orinoco River, Plants of the NYBG
from the Venezuelan Expedition 1950–1951, on bark, 19 Oct. 1950, Bassett
Maquire, R.S. Cowan & J. J. Wurdack 29223 (NY).</p><p><italic>Comments</italic>: The new name <italic>Hypocrea eucorticioides</italic> is
proposed because <italic>H. corticioides</italic> Speg. is a later homonym of <italic>H.
corticioides</italic> Berk. & Broome. The condition of the type of <italic>H.
corticioides</italic> Speg. has degraded over time, but ascospore measurements
were obtained: part-ascospores monomorphic, subglobose, (2.0–)
2.6–2.8(–4.0) × (1.9–) 2.3–2.9(–3.0) μm
(G.J. Samuels, pers. comm.). The specimens of <italic>H. eucorticioides</italic>examined from Costa Rica and Venezuela were identical to the holotype from
Argentina.</p><p>Doi (<xref ref-type="bibr" rid="ref6">1975</xref>) included
specimens in NY previously described as <italic>H. flava</italic> (from Costa Rica)
under the name <italic>H. corticioides</italic> Speg. However, the part-ascospores of
the holotype (LPS 1719 bis) illustrated by Doi
(<xref ref-type="bibr" rid="ref6">1975</xref>) in fig. 4 R and
confirmed by our study of the holotype, are monomorphic and subglobose,
differing substantially from the dimorphic part-ascospores illustrated by Doi
for specimens of <italic>H. flava,</italic> fig. 4, M, O
(<xref ref-type="bibr" rid="ref6">Doi 1975</xref>). Specimens of <italic>H.
flava</italic> were not examined in this study, but it is clear that <italic>H.
flava</italic> should be retained as a distinct species.</p></list-item><list-item><p><italic><bold>Hypocrea subsulphurea</bold></italic> Syd. in De Wildeman,<italic> Flore Bas et
Moyen-Congo</italic>: 15. 1909. Figs
<xref rid="fig13" ref-type="fig">13</xref>,
<xref rid="fig14" ref-type="fig">14</xref>.</p><p><italic>Anamorph</italic>: <italic>Trichoderma</italic> sp. [sect.
<italic>Hypocreanum</italic>].</p><p><italic>Teleomorph</italic>: Stromata effuse, extensive, surface hyphal, largest
continuous stroma 10 × 10 mm, smallest continuous stroma 2 × 2 mm,
varying in colour, usually light yellow to greyish yellow (4A5–4B5),
KOH<sup>+/–</sup>, reaction weak, slightly darkening; ostiolar canals
visible at the stroma surface, appearing golden-yellow or light orange
(4A8–6D6). Stroma surface rough, tissue immediately below the stroma
surface formed of compact to loose hyphal elements, <italic>textura
intricata</italic>. Perithecia subglobose to ellipsoidal; wall
KOH<sup>+/–</sup>, reaction variable, weak. Asci cylindrical, often
wider near the tip, (52–)59–72(–80) ×
(3.9–)4.1–5.6(–6.4) μm (n = 21); tip slightly thickened.
Part-ascospores hyaline, thick-walled, spinulose, slightly dimorphic; distal
part subglobose, (3.0–)3.4–4.0(–4.7) ×
(2.7–)2.9–3.5(–4.2) μm, L/W ratio
(0.87–)1.0–1.2(–1.4) (n = 57); proximal part subglobose to
subellipsoidal, (2.8–)3.6–4.4(–5.4) ×
(2.3–)2.9–3.6(–4.1) μm, L/W ratio
(0.90–)1.0–1.4 (–2.4) (n = 58).</p><p><italic>Anamorph</italic>: Colonies covering a 100 mm diam Petri plate with PDA in
10 d, not producing concentric rings or radial rays of mycelium; a layer of
aerial mycelium covering the entire plate; conidiophores irregularly branched,
on long hyphal elements, usually verticillium-like; phialides in whorls of
2–4, solitary, or alternating in pairs on long hyphal elements,
phialides subulate, (15.7–)22–30(–46) ×
(2.9–)3–3.7(–4.2) μm (n = 41); conidia variable in size,
ellipsoidal to subcylindrical, (5.9–)6.4–10.4(–13.5) ×
(2.7–)3.2–4.3(–4.7) μm (n = 41), truncate base not
prevalent, or when present not pronounced; no distinctive odour or pigment.
Colonies on SNA or CMD did not produce conidiophores within 10 d.</p><p><italic>Habitat</italic>: Typically found on bark with <italic>Exidia</italic> spp., also
producing new stromata over remnants of previous stroma fructifications.</p><p><italic>Known distribution</italic>: Africa and Japan.</p><p><italic>Specimen examined</italic>: <bold>Japan,</bold> Kurokami Kumamoto, Tathuta Mt.,
Research Forest at Kyushu Research Center, Forestry and Forest Products
Research Institute, old <italic>Hypocrea subsulphurea</italic> fruitbody on
<italic>Exidia</italic> sp., 14 Feb. 2002, K. Miyazaki and B.E. Overton, M141
(BPI).</p><p><italic>Comments</italic>: The type could not be located (in S) and is probably
lost. The specimen examined in this study is in poor condition and from a
different geographic location and cannot serve as neotype material. The size
of the subglobose part-ascospores and smooth-walled conidia distinguish this
species from <italic>H. microsulfurea</italic>. It is likely that <italic>H.
microsulfurea</italic> is a phylogenetic sister species of <italic>H.
subsulphurea</italic> and that globose part-ascospores and yellow extensive
stromata represent apomorphic characters. The specimen of <italic>Hypocrea
subsulphurea</italic> collected in Japan was severely degraded but discharging
ascospores. The ascospores germinated on the surface of the old specimen and
began producing additional perithecia in a thin byssoid layer. The hyphal
stromata in this specimen may be an aberration caused by <italic>in-vitro</italic>production of perithecia.</p></list-item><list-item><p><italic><bold>Hypocrea farinosa</bold></italic> Berk. & Broome, Ann. Mag. Nat. Hist.,
Ser. 2, 7: 186. 1851. Figs <xref rid="fig15" ref-type="fig">15</xref>,
<xref rid="fig16" ref-type="fig">16</xref>.</p><p><list list-type="simple"><list-item><p>≡ <italic>Protocrea farinosa</italic> (Berk. & Broome) Petch, J. Bot. 75:
219. 1937.</p></list-item></list></p><p><italic>Anamorph</italic>: <italic>Trichoderma</italic> sp. [sect.
<italic>Hypocreanum</italic>].</p><p><italic>Teleomorph</italic>: Stromata effuse, extensive, surface hyphal, largest
continuous stroma 40 × 25 mm, smallest continuous stroma 5 × 3 mm,
varying in colour, usually light yellow to light brown (4A6–6D6),
KOH<sup>+</sup>, reaction strong, with stroma becoming dark brown (6E6);
ostiolar openings visible at the stroma surface, appearing orange or light
brown (5A7–6D4). Stroma surface rough, formed of compact to loose hyphal
elements, <italic>textura intricata</italic>; below the hyphal layer is a well-defined
layer of loosely compacted pseudoparenchymatous tissue, <italic>textura
globulosa</italic>. Perithecia surrounded by a loose layer of hyphae; perithecia
generally widely spaced, compact in some regions, sometimes completely absent
near the margins or regions of extensive stroma growth. Perithecia subglobose
to subellipsoidal, (128–) 140–200(–230) μm high
(including the length of the ostiolar canal, n = 16); width of perithecia near
the base (measured from 3/4 total length of the perithecium)
(99–)110–155(–171) μm (n = 16); length of ostiolar canal
(35–)40–60(–72) μm; width of ostiolar canal from the
outer perithecial wall to the opposite internal perithecial wall
(23–)27–40(–46) μm (n =16); wall KOH<sup>+</sup>,
reaction variable. Asci cylindrical, (43–)60–90(–113)
× (2.8–) 4.0–5.6(–6.8) μm (n = 96); tip slightly
thickened. Part-ascospores hyaline, thick-walled, spinulose, dimorphic; distal
part subglobose, (2.7–)3.3–4.0(–4.8) ×
(2.3–)3–3.6(–4.7) μm, L/W ratio
(0.80–)0.98–1.2(–1.4) (n = 189); proximal part
subellipsoidal, sometimes wedge-shaped, (2.7–)3.4–4.5(–5.6)
× (2.3–)2.7–3.3(–3.9) μm, L/W ratio
(0.90–)1.1–1.5(–1.9) (n = 189).</p><p><italic>Anamorph</italic>: Colonies covering a 100 mm diam Petri plate with PDA in
10 d, not producing concentric rings or radial rays of mycelium; a layer of
aerial mycelium covering the entire plate; brown pigment near the point of
inoculation (7D7–7E7). Colonies covering a 100 mm diam Petri plate with
CMD in 10 d, producing a thin layer of effuse mycelium across the agar, with a
thin cottony layer of aerial mycelium near the agar plug and the edge of the
Petri plate; light brownish orange pigment (6C3) diffusing into the agar.
Colonies covering a 100 mm diam Petri plate with SNA in 10 d, a thin layer of
mycelium covering the agar surface, with a very thin cottony layer of aerial
mycelium near the edge of the Petri plate; pinkish white pigment (10A2)
produced near the point of inoculation. Isolates variably produce
conidiophores and conidia on all three media. A combined description of the
anamorph from all three media follows: Conidiophores produced on long hyphal
elements near the agar surface or in the aerial mycelium, irregularly
branched; phialides in whorls of (2–)3(–4); phialides subulate,
(9.5–)13–25(–36) × (1.5–)
2.4–3.0(–3.4) μm (n = 64); conidia produced terminally on
phialides, some conidia with basal abscission scar; conidia subglobose or
obovate to subellipsoidal, sometimes elongate ellipsoidal,
(3–)4.4–6.7(–11.6) ×
(1.9–)2.5–3.5(–5.0) μm (n = 79).</p><p><italic>Habitat</italic>: Fungicolous, found on lichen-covered bark,
<italic>Stereum</italic> spp., on logs inoculated with <italic>Lentinula edodes</italic>, and
in bag cultures of <italic>L. edodes</italic>.</p><p><italic>Known distribution</italic>: Japan, Europe, North America.</p><p><italic><bold>Epitype</bold></italic> (designated here): <bold>France,</bold>Pyrénées Atlantiques, Forêt Domaniale d'Oloron 64, on
bark, 30 Aug. 1997, F. Candoussau, # 513 (BPI 747356; culture G.J.S.
97-207).</p><p><italic>Other specimens examined</italic>: <bold>Canada,</bold> Ontario, Bear Island, on
<italic>Hymenochaete</italic> sp., 28 Aug. 1937, H. S. Jackson, det. R. F. Cain (BPI
631473). <bold>Japan,</bold> Morothuka Mura, anonymous mushroom farm, on log
inoculated with <italic>Lentinula edodes</italic>, 29 July 1999, K. Miyazaki, M107
(BPI). <bold>U.S.A.,</bold> Indiana, Brown County, Yellow Wood State Forest, alt.
200 m, Jackson Creek Management Trail, 39°09' N, 86°06' W, on bark, 30
Sep. 1995, G. J. Samuels (BPI 737771; culture G. J.S. 95-197); Louisiana, St.
Martinsville, on decayed wood, 24 Aug. 1890, A.B. Langlois, Flora Ludoviciana
# 2294 (BPI 631450); Maryland, Ellicot City, Paptapsco Valley State Park, on
<italic>Stereum</italic> cf. <italic>ostrea</italic>, 8 Sep. 1999, B.E. Overton, B.E.O. 99-16
(BPI; culture B.E.O. 99-16); Prince Georges County, Greenbelt Forest, on bark,
fall 1991, G. J. Samuels (BPI 1112870; culture G.J.S. 91-101); same origin, 8
Nov. 1991, S.E. Rehner (BPI 1112896); unknown mushroom farm, on wood and grain
inoculated with <italic>Lentinula edodes</italic>, 1989, sent to D. Farr at USDA/SBML
(BPI 802598; culture G.J.S. 89-139).</p><p><italic>Comments</italic>: The ascospore measurements given for <italic>Protocrea
farinosa</italic> by Rossman <italic>et al.</italic>(<xref ref-type="bibr" rid="ref15">1999</xref>), based on a
reexamination of the type, are nearly identical to those of <italic>H.
farinosa</italic> recorded in this study: distal part subglobose,
(3–)3.4–3.7(–4.6) × (2–)2.5–3(–3.3)
μm; proximal part wedge-shaped to ellipsoid, (3.2–)3.5–4.5
× 2–2.7(–3) μm. Rossman <italic>et al.</italic>(<xref ref-type="bibr" rid="ref15">1999</xref>) agreed with Doi
(<xref ref-type="bibr" rid="ref5">1972</xref>) and suggested that
<italic>P. farinosa</italic> has an acremonium-like anamorph, but noted that the
anamorph of <italic>P. farinosa</italic> described by Doi
(<xref ref-type="bibr" rid="ref5">1972</xref>) may be questionable. The
anamorph characteristics observed in this study are identical to what Doi
(<xref ref-type="bibr" rid="ref5">1972</xref>) described. The only
anomalous character is the well-defined layer of pseudoparenchymatous tissue
near the base of the perithecia, which is not consistent with the completely
hyphal stromata described in the type description of <italic>H. farinosa</italic>.
Teleomorph anatomy is variable, with some specimens of <italic>H. farinosa</italic>being more hyphal than others. In older specimens it is difficult to recognize
the pseudoparenchymatous layer near the base of the stroma in <italic>H.
farinosa</italic>. The significance of a purely hyphal stroma versus a stroma
composed of two distinct layers is unclear but it is likely that this
character has been misinterpreted because of the original condition of
specimens used to delineate <italic>H. farinosa</italic>.</p><p><italic>Hypocrea farinosa</italic> has been observed in the United States and Japan
associated with the cultivation of <italic>Lentinula edodes</italic>. Japanese
isolates of this species have been shown to be aggressive against commercial
isolates of <italic>Lentinula</italic> (Kazuhiro Miyazaki, unpublished). In the United
States, collections were made on a species of <italic>Stereum</italic> and
lichen-covered bark. <italic>Hypocrea farinosa</italic> was collected once from
<italic>Lentinula</italic> bag culture. It is likely that this species is fungicolous
in nature and is present on other fungi on logs used for production of <italic>L.
edodes</italic>; thus it can easily switch from natural substrates to the
commercial strains of <italic>L. edodes</italic>. Consequently, <italic>H. farinosa</italic>poses a greater concern to log cultivation of <italic>L. edodes</italic> than to bag
culture. An additional <italic>Hypocrea</italic> species, similar in overall
appearance to <italic>H. lactea sensu</italic> Doi, was observed associated with log
cultivation of <italic>L. edodes</italic> in Japan (Kazuhiro Miyazaki, unpublished).
Specimens of <italic>H. lactea sensu</italic> Doi were not available for direct
comparison. The epitype specimen designated here for <italic>H. farinosa</italic> from
Europe does not appear to be associated with an identifiable fungus.
Nevertheless, ITS data from this specimen are identical to those of specimens
obtained from <italic>Stereum</italic> sp., lichen-covered bark, and cultivated <italic>L.
edodes</italic>.</p></list-item><list-item><p><bold><italic>Hypocrea alcalifuscescens</italic></bold> Overton, <bold>sp. nov</bold>. MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB501057&link_type=MB">MB501057</ext-link>.
<xref rid="fig17" ref-type="fig">Fig. 17</xref>.</p><p><italic>Anamorph</italic>: <italic>Trichoderma</italic> sp. [sect.
<italic>Hypocreanum</italic>].</p><p><italic>Etymology</italic>: <italic>fuscescens</italic> (L.), turning dark in alkali; the
yellow-brown stroma of this species becomes dark reddish brown when a drop of
3 % KOH is placed on the surface.</p><p>Stromata effusa, extensa, luteo-brunnea vel olivaceo-brunnea, KOH ope
brunnescentia. Ascosporae hyalinae, crassitunicatae, spinulosae, dimorphicae;
pars distalis (3.5–)4.0–5.3(–6.7) × (3.2–)
3.7–4.5(–5.6) μm, pars proxima
(3.2–)4.5–5.8(–6.6) ×
(2.8–)3.2–4.0(–5.2) μm. Anamorphosis <italic>Trichoderma</italic>sectionis <italic>Hypocreanum</italic>. Conidia hyalina, subglobosa vel
subellipsoidea, (2.6–)4.0–7.3(–11) ×
(2.2–)2.7–4.7(–6.3) μm.</p><p>Typus: TAA(M) 181548 in BPI.</p><p><italic>Teleomorph</italic>: Stromata effuse, extensive, surface hyphal; largest
continuous stroma 25 × 10 mm, varying in colour, usually yellow-brown to
olive-brown (5E8–4E7), KOH<sup>+</sup>, darkening, reddish brown (8E8);
ostiolar canals visible at the stroma surface, appearing brown (6E8), stroma
surface with furrows; tissue immediately below the stroma surface formed of
compact hyphal elements, textura intricata. Perithecia subglobose to
ellipsoidal; wall KOH<sup>+/–</sup>, reaction variable, weak. Asci
cylindrical, often wider near the tip, (78–)90–112(–121)
× (4.0–)4.4-5.7(–6.4) μm (n = 30); tip slightly
thickened. Part-ascospores hyaline, thick-walled, spinulose, dimorphic; distal
part subglobose, sometimes obovate, (3.5–)4.0–5.3(–6.7)
× (3.2–)3.7–4.5(–5.6) μm, L/W ratio
(0.87–)1.0–1.3(–1.5) (n = 65); proximal part subglobose to
oblong ellipsoidal, sometimes wedge-shaped,
(3.2–)4.5–5.8(–6.6) ×
(2.8–)3.2–4.0(–5.2) μm, L/W ratio
(0.90–)1.2–1.6(–2.1) (n = 65).</p><p><italic>Anamorph</italic>: Colonies covering a 100 mm diam Petri plate with PDA in
10 d, not producing concentric rings or radial rays of mycelium; a layer of
aerial mycelium covering the entire plate; conidiophores irregularly branched,
on long hyphal elements, usually verticillium-like; phialides in whorls of
4–6(–8), rarely solitary, or alternating in pairs; phialides
subulate, (6.7–)13–24(–30.3) ×
(1.6–)2.3–3.0(–3.5) μm (n = 38); conidia variable in
size, subglobose to obovate, subellipsoidal, rarely subcylindrical,
(2.6–)4.0–7.3(–11) ×
(2.2–)2.7–4.7(–6.3) μm (n = 63), infrequently with an
indistinct flat edge; no distinctive odour, brownish orange pigment (6C4)
produced near the point of inoculation. Colonies on SNA or CMD did not produce
conidiophores within 10 d.</p><p><italic>Habitat</italic>: Possibly fungicolous, found on leaf litter with
<italic>Piloderma/Amauroderma</italic>, also known from bark of <italic>Liriodendron
tulipifera</italic> without visible evidence of another fungus present.</p><p><italic>Known distribution</italic>: Eastern Europe and North America, apparently
not common.</p><p><italic>Holotype</italic>: <bold>Estonia,</bold> on leaf litter and
<italic>Piloderma/Amauroderma</italic>, 13 Sep. 2000, U. Kõljalg, TAA(M) 181548
(BPI, ex-type culture TFC 2000-36).</p><p><italic>Other specimen examined</italic>: <bold>U.S.A.,</bold> Delaware, Rockland, on
bark of <italic>Liriodendron tulipifera,</italic> 7 July 1890 (BPI 631474; herb. J. B.
Ellis).</p></list-item><list-item><p><italic><bold>Hypocrea parmastoi</bold></italic> Overton, <bold>sp. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB501058&link_type=MB">MB501058</ext-link>.
<xref rid="fig18" ref-type="fig">Fig. 18</xref>.</p><p><italic>Anamorph</italic>: <italic>Trichoderma</italic> sp. [sect.
<italic>Hypocreanum</italic>].</p><p><italic>Etymology</italic>: This species is named after Dr E. Parmasto in
recognition of his significant mycological contributions, and in appreciation
of his assistance in identifying polypores in this study.</p><p>Stromata effusa, extensa, violaceo-brunnea vel griseo-purpurea,. Ascosporae
hyalinae, crassitunicatae, spinulosae, dimorphicae; pars distalis ascosporarum
subglobosa vel obovata, (2.7–)3.8–5.2(–6.7) ×
(2.7–)3.5–4.5(–5.6) μm, pars proxima sublgoobsa vel
ellipsoidea, (3.2–)4.2–5.7(–6.6) ×
(2.4–)3.0–4.0(–5.2) μm. Anamorphosis <italic>Trichoderma</italic>sectionis <italic>Hypocreanum.</italic> Conidia hyalina, subglobosa vel subcylindrica,
(3.0–)3.4–5.0(–7.8) ×
(2.2–)2.5–3.1(–3.6) μm.</p><p><italic>Teleomorph</italic>: Stromata effuse, extensive, surface hyphal, largest
continuous stroma 20 × 10 mm, varying in colour, usually violet-brown to
greyish ruby (10E7–12DE7), KOH<sup>–</sup>; ostiolar canals
visible at the stroma surface, greyish brown (10F3); stroma surface rugulose,
tissue immediately below the stroma surface formed of compact to loose hyphal
elements, textura intricata. Perithecia subglobose to ellipsoidal; wall
KOH<sup>+/–</sup>, reaction variable, weak. Asci cylindrical, often
wider near the tip, (78–)89–112(–121) ×
(3.9–)4.4–5.6(–6.4) μm (n = 30); tip slightly
thickened.</p><p>Part-ascospores hyaline, thick-walled, spinulose, dimorphic; distal part
subglobose, sometimes obovate, (2.7–)3.8–5.2(–6.7) ×
(2.7–)3.5–4.5(–5.6) μm, L/W ratio
(0.75–)1.0–1.3(–1.4) (n = 77); proximal part, subglobose to
ellipsoidal, (3.2–)4.2–5.7(–6.6) × (2.4–)
3.0–4.0(–5.2) μm, L/W ratio
(0.90–)1.2–1.6(–2.1) (n = 77).</p><p><italic>Anamorph</italic>: Colonies covering a 100 mm diam Petri plate with PDA in
10 d, producing radial rays of mycelium and a layer of aerial mycelium near
the agar plug and the edge of the plate; conidiophores irregularly branched,
on long hyphal elements, usually verticillium-like; phialides in whorls of
2–4(–10), sometimes solitary or in pairs forming a fork shape;
phialides subulate, (12–)16–29(–33.3) ×
(1.6–)2.1–3.0(–3.4) μm (n = 34); conidia variable in
shape, subglobose to obovate, or subellipsoidal to subcylindrical,
(3.0–)3.4–5.0(–7.8) ×
(2.2–)2.5–3.1(–3.6) μm (n = 38), infrequently with an
flat base, which is then not conspicuous; no distinctive odour; dark rose
pigment (11A3) produced near the point of inoculation. Colonies covering a 100
mm diam Petri plate with CMD in 10 d, producing a layer of aerial mycelium
near the agar plug and the edge of the plate; conidiophores irregularly
branched, on long hyphal elements, usually verticillium-like; phialides in
whorls of 2–4(–10), sometimes solitary or in pairs; phialides
subulate, with the same measurements as on PDA; no distinctive odour; reddish
white pigment (11A2) produced near the point of inoculation. Colonies on SNA
did not produce any conidiophores within 10 d.</p><p><italic>Habitat</italic>: Lignicolous, on <italic>Alnus incana</italic> and
<italic>Castanea</italic> sp., possibly <italic>Quercus</italic> spp.</p><p><italic>Known distribution</italic>: Eastern Europe and France, apparently not
common.</p><p><italic>Holotype</italic>: <bold>Estonia,</bold> Võru Commune, Võrumaa
County, 57°47' N, 27°9' E, on <italic>Alnus incana</italic> (fallen trunk),
Kütiorg, 3 Oct. 1997, I. Parmasto, TAA(M) 169055 (BPI; ex-type culture
TFC 97-143).</p><p><italic>Other specimen examined</italic>: <bold>France,</bold> St. Gaudens 31, Haute
Garonne, Arboretum De Cudeilhac, on <italic>Castanea</italic> sp., possibly
<italic>Quercus</italic> sp., 1 Nov. 1994, Françoise Candoussau (BPI
737853).</p></list-item></list></p><p><fig position="float" id="fig6"><label>Fig. 6.</label><caption><p>A–C. <italic>H. sulphurea</italic>. A. Stroma with KOH reaction, BPI 737764;
bar = 1 mm. B. Stroma with byssoid margin, BPI 737752; bar = 1 mm. C. Stroma
with KOH reaction, TNS-F-190169; bar = 1 mm.</p></caption><graphic xlink:href="39fig6"></graphic></fig></p><p><fig position="float" id="fig7"><label>Fig. 7.</label><caption><p>A–F. <italic>H. sulphurea</italic>. A. Section of stroma showing ostiolar
papilla; bar = 20 μm. B. Section of stroma showing <italic>textura
globulosa</italic> to <italic>t. angularis</italic> with <italic>t. intricata</italic> below
perithecium; bar = 20 μm. C. Asci with ascospores; bar = 20 μm;
A–C. BPI 737752. D. Section showing <italic>t. angularis</italic> near surface,
B.E.O. 98-44. E, F. TNS-F-190169, <italic>H. sulphurea</italic> f.
<italic>macrospora</italic>; E. Section through ostiole; F. asci; bars = 20 μm. G.
Smooth stroma surface, BPI 747764, <italic>H. sulphurea</italic> f.
<italic>sulphurea</italic>; bar = 1 mm.</p></caption><graphic xlink:href="39fig7"></graphic></fig></p><p><fig position="float" id="fig8"><label>Fig. 8.</label><caption><p>A–F. <italic>H. sulphurea</italic>. A. Conidia of variable size, on PDA; bar
= 20 μm. B–C. Conidiophores, on PDA; bars = 20 μm; A–C.
B.E.O. 98-44. D-F. Conidiophores and conidia from G.J.S. 95-140 on PDA; bars =
20 μm.</p></caption><graphic xlink:href="39fig8"></graphic></fig></p><p><fig position="float" id="fig9"><label>Fig. 9.</label><caption><p>A–F. <italic>H. victoriensis</italic>. A. Stroma with discharged ascospores,
BPI 747361; bar = 1 mm. B. Stroma showing KOH reaction, BPI 747363; bar = 1
mm. C. Section of Stroma, showing ostiolar papillae; bar = 40 μm. D, F.
Section of stroma showing <italic>t. globulosa</italic> to <italic>t. angularis</italic>; bar
= 20 μm. E. Asci with spinulose ascospores; bar = 20 μm; C–E. BPI
747362.</p></caption><graphic xlink:href="39fig9"></graphic></fig></p><p><fig position="float" id="fig10"><label>Fig. 10.</label><caption><p>A–E. <italic>H. victoriensis</italic>. A–C. Irregular verticillium-like
conidiophore branching pattern; bars = 20 μm. D. Conidiophore, G. J. S.
99-201; bar = 20 μm. E. Conidia; bar = 20 μm; A–C, E. G.J.S.
99-200.</p></caption><graphic xlink:href="39fig10"></graphic></fig></p><p><fig position="float" id="fig11"><label>Fig. 11.</label><caption><p>A–F. <italic>H. eucorticioides</italic>. A. Effuse stromata with irregular
margins, NY 29223; bar = 1 mm. B. Effuse stromata with irregular margins
showing reaction in KOH, BPI 747358; bar = 1 mm. C. Asci with subglobose
part-ascospores, BPI 747358; bar = 20 μm. D. Asci with subglobose
part-ascospores, NY 29223; bar = 20 μm. E–F. Section of stroma
showing <italic>t. globulosa</italic> to <italic>t. angularis</italic> tissue, BPI 747358; bar
= 20 μm.</p></caption><graphic xlink:href="39fig11"></graphic></fig></p><p><fig position="float" id="fig12"><label>Fig. 12.</label><caption><p>A–D. <italic>H. eucorticioides</italic>, G.J.S. 99-61. A. Irregular
verticillium-like branching pattern; bar = 40 μm. B–D. Phialides with
developing conidia; bar = 20 μm.</p></caption><graphic xlink:href="39fig12"></graphic></fig></p><p><fig position="float" id="fig13"><label>Fig. 13.</label><caption><p>A–D. <italic>H. subsulphurea</italic>, M 141. A–B. Perithecia forming
on old stromata; bar = 1 mm. C. Asci and ascospores from old stroma; bar = 20
μm. D. Asci and ascospores from perithecia developing on old stroma
surface; bar = 20 μm. Note: repeated attempts to section perithecia in old
and developing stromata have failed.</p></caption><graphic xlink:href="39fig13"></graphic></fig></p><p><fig position="float" id="fig14"><label>Fig. 14.</label><caption><p>A–D. <italic>H. subsulphurea</italic>, M. 141. A–B. Irregular
verticillium-like branching pattern, on PDA; bar = 40 μm. C. Phialides with
developing conidia, on PDA; bar = 20 μm. D. Conidia, variable in size, on
PDA; bar = 20 μm.</p></caption><graphic xlink:href="39fig14"></graphic></fig></p><p><fig position="float" id="fig15"><label>Fig. 15.</label><caption><p>A–D. <italic>H. farinosa</italic>. A. Effuse extensive stroma, BPI 802598. B.
Effuse extensive stroma, BPI 747356. C. Colour variation in stroma, BPI
737771. D. KOH reaction, BPI 802598. bars = 1 mm.</p></caption><graphic xlink:href="39fig15"></graphic></fig></p><p><fig position="float" id="fig16"><label>Fig. 16.</label><caption><p>A–H. <italic>H. farinosa</italic>. A. Section of stroma; note the layer of
<italic>t. intricata</italic> near the stroma surface and loose layer of
pseudoparenchymatous tissue near the base of the perithecium; bar = 40 μm.
B. asci and part-ascospores, both of BPI 802598, bar = 20 μm. C. Section of
stroma; bar = 40 μm. D–E. <italic>Textura intricata</italic> near stroma
surface and <italic>t. globulosa</italic> to <italic>t. angularis</italic> below perithecium,
notably different from that shown in A; bar = 20 μm. C–E. BPI 112870.
F–G. Conidiophores and developing conidia, on PDA; bars = 20 μm. H.
Conidia on PDA; bar = 40 μm; F–H. G.J.S. 89-139.</p></caption><graphic xlink:href="39fig16"></graphic></fig></p><p><fig position="float" id="fig17"><label>Fig. 17.</label><caption><p>A–F. <italic>H. alcalifuscescens</italic>, TAA 181584. A. Stroma showing
reaction in KOH; bar = 1 mm. B. Section of stroma showing compact <italic>t.
intricata</italic> and KOH-positive layer of stroma tissue; bar = 20 μm. C.
Asci and ascospores; bar = 20 μm. D. Conidia; bar = 20 μm. E–F.
Conidiophore branching pattern; bars = 40 μm, and 20 μm, respectively;
D–F. TFC 181584. Note: in spite of several attempts, no illustrative
sections of perithecia were obtained.</p></caption><graphic xlink:href="39fig17"></graphic></fig></p><p><fig position="float" id="fig18"><label>Fig. 18.</label><caption><p>A–F. <italic>H. parmastoi</italic>. A–B. Stroma; bars = 1 mm and 20
μm, respectively. C. Asci and ascospores; bar = 20 μm; A–C. TAA
169055. D, F. Conidiophores on PDA; bar = 20 μm. E. Conidiophores on CMD;
bar = 40 μm; D–F. TFC 97-143. Note: in spite of several attempts, no
illustrative sections of perithecia were obtained.</p></caption><graphic xlink:href="39fig18"></graphic></fig></p></sec><sec><title>KEY TO THE SPECIES TREATED</title><p><list list-type="simple"><list-item><p>1. Occurring on <italic>Exidia</italic> spp.; stromata yellow, effuse,
extensive............................................................................
2</p></list-item><list-item><p>1. Not occurring on <italic>Exidia</italic> spp.; stromata variable in colour,
discrete to extensive................................................ 3</p></list-item><list-item><p>2. Part-ascospores dimorphic, hyaline, thick-walled, spinulose; distal part
obovate, sometimes subellipsoidal, (4.2–)5.2–6.6(–7.6)
× (4.2–)5.3–7.1(–8.3) μm, proximal part
ellipsoidal, sometimes subcylindrical, (4.4–)5.5–6.9(–8.5)
× (2.7–)3.9–5.1(–6.6)
μm.................................. 1. <italic><bold>H. sulphurea</bold></italic></p></list-item><list-item><p>2. Part-ascospores monomorphic, hyaline, thick-walled, spinulose,
subglobose to sub-ellipsoidal, (2.8–)3.6–4.4(–5.4) ×
(2.3–)2.9–3.6(–4.1)
μm.................................................... 5. <italic><bold>H.
subsulphurea</bold></italic></p></list-item><list-item><p>3. Stromata hyphal or sometimes with a thin layer of pseudoparenchymatous
tissue below the
perithecia...........................................................................................................................................
4</p></list-item><list-item><p>3. Stroma tissue pseudoparenchymatous, no hyphal layer; stromata yellow to
yellow-brown, effuse or
subpulvinate............................................................................................................................
6</p></list-item><list-item><p>4. Stromata composed of a loose layer of hyphae, violet-brown;
part-ascospores dimorphic, hyaline, thick-walled, spinulose; distal part
subglobose, sometimes obovate, (3.5–)4.0–5.3(–6.7) ×
(3.2–)3.7–4.5(–5.6) μm, proximal part subglobose to
subellipsoidal, (3.2–)4.5–5.8(–6.6) ×
(2.8–)3.2–4.0(–5.2)
μm............................................................................
8. <italic><bold>H. parmastoi</bold></italic></p></list-item><list-item><p>4. Stromata either composed of compact hyphae, or with both a hyphal layer
near the surface and a pseudoparenchymatous layer below the
perithecia....................................................................
5</p></list-item><list-item><p>5. Stromata composed of tightly packed hyphae, olive-brown at the surface;
part-ascospores dimorphic, hyaline, thick-walled, spinulose; distal part
subglobose, sometimes obovate, (3.5–)4.0–5.3(–6.7) ×
(3.2–)3.7–4.5(–5.6) μm, proximal part subglobose to
subellipsoidal, (3.2–)4.5–5.8(–6.6) ×
(2.8–)3.2–4.0 (–5.2)
μm................................................................ 7.
<italic><bold>H. alcalifuscescens</bold></italic></p></list-item><list-item><p>5. Stromata composed of two layers, a hyphal layer near the surface, and a
pseudoparenchymatous layer near the base of the perithecia, light yellow to
light brown; part-ascospores dimorphic, hyaline, thick-walled, spinulose,
distal part subglobose, (2.7–)3.3–4.0(–4.8) ×
(2.3–)3–3.6(–4.7) μm, proximal part subellipsoidal,
sometimes wedge-shaped, (2.7–)3.4–4.5(–5.6) ×
(2.3–)2.7–3.3 (–3.9)
μm..................................................... 6. <italic><bold>H.
farinosa</bold></italic></p></list-item><list-item><p>6. Part-ascospores monomorphic, subglobose, spinulose,
(2.6–)2.8–3.5(–3.9) ×
(2.4–)2.6–3.2(–3.7) μm; stromata yellow, effuse,
extensive................................................ 4. <italic><bold>H.
eucorticioides</bold></italic></p></list-item><list-item><p>6. Part-ascospores dimorphic; stromata effuse to subpulvinate, yellow to
yellowish
brown..................................................................................................................................................................
7</p></list-item><list-item><p>7. Part-ascospores dimorphic, hyaline, thick-walled, spinulose; distal part
subellipsoidal, sometimes obovate, (4.8–)5.6–6.8(–7.4)
× (3.9–)4.5–5.5(–5.9) μm, proximal part
ellipsoidal, sometimes wedge-shaped, (4.7–)5.8–7.4(–8.6)
× (3.6–)4.1–5.1(–6.1)
μm.............................. 3. <italic><bold>H. victoriensis</bold></italic></p></list-item><list-item><p>7. Part-ascospores dimorphic, distal part subglobose to obovate conical,
(4.5–)4.7–5.1 (–5.4) ×
(3.6–)3.8–4.2(–4.3) μm; proximal part tending to be
oblong or ellipsoidal, narrow, (4.2–)4.7–5.6(–6.0) ×
(3.6–)3.5–3.7(–3.8) μm; known from Africa and New
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