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<record><TEI><teiHeader><fileDesc><titleStmt><title xml:lang="en">Systematics of <italic>Hypocrea citrina</italic> and related taxa</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><author><name sortKey="Jaklitsch, Walter M" sort="Jaklitsch, Walter M" uniqKey="Jaklitsch W" first="Walter M." last="Jaklitsch">Walter M. Jaklitsch</name><affiliation><nlm:aff id="aff3"><italic>TU Vienna, Institute of Chemical Engineering, Division Applied Biochemistry and Gene Technology, x Getreidemarkt 9/E1665, A-1060 Vienna, Austria.</italic></nlm:aff></affiliation></author></titleStmt><publicationStmt><idno type="wicri:source">PMC</idno><idno type="pmid">18490988</idno><idno type="pmc">2104732</idno><idno type="url">http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2104732</idno><idno type="RBID">PMC:2104732</idno><idno type="doi">10.3114/sim.2006.56.01</idno><date when="2006">2006</date><idno type="wicri:Area/Pmc/Corpus">000101</idno><idno type="wicri:explorRef" wicri:stream="Pmc" wicri:step="Corpus" wicri:corpus="PMC">000101</idno></publicationStmt><sourceDesc><biblStruct><analytic><title xml:lang="en" level="a" type="main">Systematics of <italic>Hypocrea citrina</italic> and related taxa</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><author><name sortKey="Jaklitsch, Walter M" sort="Jaklitsch, Walter M" uniqKey="Jaklitsch W" first="Walter M." last="Jaklitsch">Walter M. Jaklitsch</name><affiliation><nlm:aff id="aff3"><italic>TU Vienna, Institute of Chemical Engineering, Division Applied Biochemistry and Gene Technology, x Getreidemarkt 9/E1665, A-1060 Vienna, Austria.</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 three
genomic regions – 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 translation elongation factor
(<italic>tef1</italic>) – were used to investigate the systematics of
<italic>Hypocrea citrina</italic> and related species. A neotype specimen is
designated for <italic>H. citrina</italic> that conforms to Persoon's description of a
yellow effuse fungus occurring on leaf litter. Historical information and
results obtained in this study provide the foundation for selection of a
lectotype specimen from Fries's herbarium for <italic>H. lactea</italic>. The results
indicate that (1) <italic>Hypocrea citrina</italic> and <italic>H. pulvinata</italic> are
distinct species; (2) <italic>H. lactea sensu</italic> Fries is a synonym of the older
name <italic>H. citrina</italic>; (3) <italic>H. pulvinata, H. protopulvinata</italic>, and
<italic>H. americana</italic> are phylogenetically distinct species that form a
well-supported polyporicolous clade; (4) <italic>H. citrina</italic> is situated in a
clade closely related to <italic>H. pulvinata</italic>; and (5) <italic>H.
microcitrina</italic> and <italic>H. pseudostraminea</italic> reside in a highly supported
clade phylogenetically distinct from <italic>H. citrina. Hypocrea protopulvinata,
H. microcitrina, H. megalocitrina, H. pseudostraminea</italic>, and a new species,
<italic>H. aurantiistroma</italic>, are reported and described from North America.
Variation in <italic>rpb2</italic> and <italic>tef1</italic> gene sequences suggests
geographical subgroupings between European and North American isolates of
<italic>H. pulvinata</italic>. The phylogenies inferred from ITS, <italic>rpb2</italic>, and
<italic>tef1</italic> gene sequences are concordant. <italic>Hypocrea citrina</italic> var.
<italic>americana</italic> is elevated to species status, <italic>Hypocrea
americana</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><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">18490988</article-id><article-id pub-id-type="pmc">2104732</article-id><article-id pub-id-type="publisher-id">0001</article-id><article-id pub-id-type="doi">10.3114/sim.2006.56.01</article-id><article-categories><subj-group subj-group-type="heading"><subject>Articles</subject></subj-group></article-categories><title-group><article-title>Systematics of <italic>Hypocrea citrina</italic> and related taxa</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 contrib-type="author"><name><surname>Jaklitsch</surname><given-names>Walter M.</given-names></name><xref ref-type="aff" rid="aff3">3</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><aff id="aff3"><label>3</label><italic>TU Vienna, Institute of Chemical Engineering, Division Applied Biochemistry and Gene Technology, x Getreidemarkt 9/E1665, A-1060 Vienna, Austria.</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>1</fpage><lpage>38</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="1.pdf"></self-uri><abstract><p>Morphological studies and phylogenetic analyses of DNA sequences from three
genomic regions – 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 translation elongation factor
(<italic>tef1</italic>) – were used to investigate the systematics of
<italic>Hypocrea citrina</italic> and related species. A neotype specimen is
designated for <italic>H. citrina</italic> that conforms to Persoon's description of a
yellow effuse fungus occurring on leaf litter. Historical information and
results obtained in this study provide the foundation for selection of a
lectotype specimen from Fries's herbarium for <italic>H. lactea</italic>. The results
indicate that (1) <italic>Hypocrea citrina</italic> and <italic>H. pulvinata</italic> are
distinct species; (2) <italic>H. lactea sensu</italic> Fries is a synonym of the older
name <italic>H. citrina</italic>; (3) <italic>H. pulvinata, H. protopulvinata</italic>, and
<italic>H. americana</italic> are phylogenetically distinct species that form a
well-supported polyporicolous clade; (4) <italic>H. citrina</italic> is situated in a
clade closely related to <italic>H. pulvinata</italic>; and (5) <italic>H.
microcitrina</italic> and <italic>H. pseudostraminea</italic> reside in a highly supported
clade phylogenetically distinct from <italic>H. citrina. Hypocrea protopulvinata,
H. microcitrina, H. megalocitrina, H. pseudostraminea</italic>, and a new species,
<italic>H. aurantiistroma</italic>, are reported and described from North America.
Variation in <italic>rpb2</italic> and <italic>tef1</italic> gene sequences suggests
geographical subgroupings between European and North American isolates of
<italic>H. pulvinata</italic>. The phylogenies inferred from ITS, <italic>rpb2</italic>, and
<italic>tef1</italic> gene sequences are concordant. <italic>Hypocrea citrina</italic> var.
<italic>americana</italic> is elevated to species status, <italic>Hypocrea
americana</italic>.</p></abstract><kwd-group><kwd>Ascomycetes</kwd><kwd><italic>Hypocrea citrina</italic></kwd><kwd><italic>Hypocrea lactea</italic></kwd><kwd><italic>Hypocrea pulvinata</italic></kwd><kwd><italic>Hypocreales</italic></kwd><kwd><italic>Hypocreanum</italic></kwd><kwd>ITS</kwd><kwd>systematics</kwd><kwd><italic>rpb2</italic> gene sequences</kwd><kwd><italic>tef1</italic> gene sequences</kwd><kwd>Subsection <italic>Citrinae</italic></kwd><kwd>Subsection <italic>Pulvinatae</italic></kwd><kwd><italic>Trichoderma</italic></kwd></kwd-group></article-meta><notes><fn-group><fn><p><bold>Taxonomic novelties:</bold> A neotype is designated for <italic>Hypocrea
citrina</italic> (Pers: Fr.) Fr. and a lectotype for <italic>Hypocrea lactea</italic>(Fr.: Fr.) Fr.; <italic>H.</italic> <italic>pulvinata</italic> Fuckel is lectotypified;
<italic>Hypocrea aurantiistroma</italic> Overton, sp. nov., <italic>Hypocrea
americana,</italic> stat. nov.</p></fn></fn-group></notes></front><body><sec><title>INTRODUCTION</title><p>Among the most common and conspicuous members of the genus
<italic>Hypocrea</italic> (<italic>Hypocreales, Hypocreaceae</italic>) are species that form
extensively effused, lightly or brightly coloured stromata on leaf litter,
soil, wood, and bracket fungi. Names that have been applied to these fungi
include <italic>H. citrina</italic> (Pers.: Fr.) Fr., <italic>H. lactea</italic> (Fr.: Fr.)
Fr., <italic>H. pulvinata</italic> Fuckel, <italic>H. fungicola</italic> P. Karst., and <italic>H.
karsteniana</italic> Niessl. Confusion surrounds the application of the name
<italic>H. citrina</italic> (Pers.: Fr.) Fr., and phylogenetically related species
have not been determined. The stroma of <italic>H. citrina</italic> is extensively
effuse, typically yellow and asci contain eight 2-celled uniseriate
ascospores.</p><p>Doi (<xref ref-type="bibr" rid="ref9">1972</xref>) subdivided
<italic>Hypocrea</italic> into subgenera, sections, subsections, and series based on
stroma characters. He recognized <italic>H. pulvinata, H. citrina</italic>, and <italic>H.
lactea</italic> as distinct species and placed them in <italic>Hypocrea</italic> subgen.
<italic>Hypocrea</italic> sect. <italic>Homalocrea</italic> Yoshim. Doi. He included <italic>H.
citrina</italic> and <italic>H. lactea</italic> in subsect. <italic>Citrinae</italic> Yoshim. Doi,
while <italic>H. pulvinata</italic> was placed in subsect <italic>Pulvinatae</italic> Yoshim.
Doi. The anamorph of <italic>H. citrina</italic> has acremonium- or verticillium-like
conidiophores with hyaline conidia. Bissett
(<xref ref-type="bibr" rid="ref1">1991a</xref>,
<xref ref-type="bibr" rid="ref2">b</xref>) assigned acremonium- or
verticillium-like condiophores to <italic>Trichoderma</italic> sect<italic>.
Hypocreanum</italic> Bissett. Fourteen species of <italic>Hypocrea</italic> Fr., with
stromata similar to that of <italic>H. citrina</italic>, or considered as its synonym,
and anamorphs assignable to <italic>Trichoderma</italic> sect. <italic>Hypocreanum</italic>are newly described or redescribed, type specimens examined, and, where
possible, phylogenetic relationships to <italic>H. citrina</italic> established.</p><sec><title>Nomenclature and typification</title><p>The name <italic>H. citrina</italic> has been applied to two different species, one
polyporicolous and one lignicolous or terrestrial. Rifai & Webster
(<xref ref-type="bibr" rid="ref28">1966</xref>) considered the
terrestrial/lignicolous form of <italic>H. citrina</italic> to be a synonym of another
terrestrial species, <italic>H. lactea</italic>. Canham
(<xref ref-type="bibr" rid="ref3">1969</xref>) used the name <italic>H.
citrina,</italic> with <italic>H. pulvinata</italic> as a synonym, for the polyporicolous
form. Perspectives on these differing conclusions can be found in Rifai &
Webster (<xref ref-type="bibr" rid="ref28">1966</xref>) and Canham
(<xref ref-type="bibr" rid="ref3">1969</xref>). Application of the
current International Code of Botanical Nomenclature (ICBN:
<xref ref-type="bibr" rid="ref16">Greuter <italic>et al.</italic> 2000</xref>)
for the selection of type material provides the mechanism for stabilizing the
application of the names, <italic>H. citrina, H. lactea</italic>, and <italic>H.
pulvinata</italic>.</p><p>Persoon (<xref ref-type="bibr" rid="ref27">1796</xref>) described a
yellow terrestrial fungus on leaf litter as <italic>Sphaeria citrina</italic> Pers.
Fries (<xref ref-type="bibr" rid="ref11">1816</xref>) described
<italic>Sphaeria lactea</italic> Fr. as a white effuse fungus with dark ostioles that
occurs on bark and soil. Fries
(<xref ref-type="bibr" rid="ref13">1823</xref>) expanded Persoon's
concept of <italic>Sphaeria citrina</italic> Pers.: Fr. to include a polyporicolous
element, listing <italic>Scleromyceti Sueciae</italic> 31 (<italic>SS</italic> 31) as
representative material. Fries
(<xref ref-type="bibr" rid="ref14">1849</xref>) described the genus
<italic>Hypocrea</italic> (formerly a section of <italic>Sphaeria</italic>) and transferred
<italic>S. citrina</italic> and <italic>S. lactea</italic> into <italic>Hypocrea</italic>. Persoon's
(<xref ref-type="bibr" rid="ref27">1796</xref>) original protologue
clearly emphasized a yellow effuse terrestrial species that occurs on leaf
litter. Fries's expanded species concept caused confusion. He explicitly
ascribed the species, however, to Persoon in the Index to the same volume and
to volume 3 of his <italic>Systema mycologicum.</italic> Dingley
(<xref ref-type="bibr" rid="ref7">1955</xref>) and Canham
(<xref ref-type="bibr" rid="ref3">1969</xref>) followed Fries's
(<xref ref-type="bibr" rid="ref13">1823</xref>) concept of <italic>H.
citrina</italic> to be a fungus that occurs on polypores, even though,
historically, others such as Karsten
(<xref ref-type="bibr" rid="ref17">1873</xref>), Niessl
(<xref ref-type="bibr" rid="ref22">1883</xref>) and Saccardo
(<xref ref-type="bibr" rid="ref29">1883</xref>) had recognized the
fungicolous element included by Fries as taxonomically distinct from <italic>S.
citrina</italic>. To account for the polyporicolous element that Fries
(<xref ref-type="bibr" rid="ref13">1823</xref>) ascribed to <italic>S.
citrina</italic>, Karsten (<xref ref-type="bibr" rid="ref17">1873</xref>)
published the new name, <italic>Hypocrea citrina</italic> * <italic>fungicola</italic> P.
Karst., without designating a formal rank. Niessl
(<xref ref-type="bibr" rid="ref22">1883</xref>) raised Karsten's taxon
to species rank, <italic>H. karsteniana</italic> Niessl. Only months later, Saccardo
(<xref ref-type="bibr" rid="ref29">1883</xref>), apparently unaware of
Niessl's work, also elevated Karsten's taxon to species rank as <italic>H.
fungicola</italic> (P. Karst.) Sacc. Karsten, Niessl, and Saccardo did not
recognize <italic>H. pulvinata</italic> as a synonym, although it had previously been
described from a polypore (<xref ref-type="bibr" rid="ref15">Fuckel
1870</xref>). Seaver
(<xref ref-type="bibr" rid="ref30">1910</xref>) listed <italic>H.
karsteniana</italic> and <italic>H. fungicola</italic> as questionable synonyms of <italic>H.
citrina</italic>. Weese (<xref ref-type="bibr" rid="ref33">1927</xref>)
synonymized <italic>H. karsteniana</italic> and <italic>H. fungicola</italic> with the oldest
available name, <italic>H. pulvinata</italic>. The synonymy proposed by Weese
(<xref ref-type="bibr" rid="ref33">1927</xref>), which has never been
disputed, serves as the basis for the application of the name <italic>H.
pulvinata</italic> to the polyporicolous species.</p><p>There is no specimen of <italic>S. citrina</italic> in Persoon's herbarium (L), and
collections that can be directly linked to Persoon cannot be located. Dingley
(<xref ref-type="bibr" rid="ref7">1955</xref>) took Fries's <italic>SS</italic>31 (UPS), on a polypore, to be the type collection but did not formally
designate the specimen as lectotype. Canham
(<xref ref-type="bibr" rid="ref3">1969</xref>) formally designated
<italic>SS</italic> 31 (UPS) to be the lectotype of <italic>S. citrina</italic>. Canham
(<xref ref-type="bibr" rid="ref3">1969</xref>) described <italic>H.
citrina</italic> var. <italic>americana</italic> Canham, with globose part-ascospores from
polypores in North America. Fries
(<xref ref-type="bibr" rid="ref13">1823</xref>) adopted Persoon's
epithet, thereby sanctioning it, and the name is therefore now cited as
<italic>Hypocrea citrina</italic> (Pers.: Fr.) Fr. Article 7.8 ICBN allows the
selection of neotypes among material cited by the sanctioning authority. In
this case, the taxonomic circumscription of the name and its type specimen
clearly comes from Persoon and Fries's species concept was confused. Fries's
interpreation deviates from the protologue and thus we reject Canham's
(<xref ref-type="bibr" rid="ref3">1969</xref>) lectotypification.</p><p>Rifai & Webster (<xref ref-type="bibr" rid="ref28">1966</xref>)
rejected Dingley's (<xref ref-type="bibr" rid="ref7">1955</xref>) [and
Canham's (<xref ref-type="bibr" rid="ref3">1969</xref>)] concept of
<italic>H. citrina</italic> as being fungicolous, because it did not agree with the
protologue of <italic>S. citrina.</italic> Based on the argument proposed by Rifai
& Webster (<xref ref-type="bibr" rid="ref28">1966</xref>), it is
evident that they wished to preserve the distinction between the fungicolous
and terrestrial/lignicolous species in order to retain the name <italic>H.
pulvinata</italic> for the fungicolous species. Their argument emphasized the
original protologues of <italic>H. citrina</italic> and <italic>H. pulvinata</italic>. Rifai
& Webster (<xref ref-type="bibr" rid="ref28">1966</xref>)
designated Persoon's original description of <italic>S. citrina</italic> as the
lectotype in contravention to Arts. 8.1 and 9.2 of the ICBN that state a
lectotype must be a specimen, or, at least, an illustration. By designating a
litter-inhabiting specimen as neotype, we intend to stabilize the application
of the name <italic>H. citrina</italic>.</p><p>Fries (<xref ref-type="bibr" rid="ref11">1816</xref>) described
<italic>Sphaeria lactea</italic> as “... latissime effusa subcarnosa lactea
nuda, ostiolis punctiformibus immersis prominulisque”. He reported that
the new species grows on fallen logs and on naked soil in moist places and
that the white colour does not change over time. He did not cite a specific
collection or collecting locality. There are two specimens in Fries's
herbarium (UPS) that can possibly be considered for neotypifying <italic>S.
lactea</italic>, both of which were collected by Fries in “Småland,
Femsjö”; one is identified as <italic>H. lactea</italic> and the other as
<italic>Sphaeria lactea</italic>, but neither specimen has a date of collection. The
specimen of <italic>H. lactea</italic> comprises an effused, yellow <italic>Hypocrea</italic>stroma that completely covers the leaf of a dicotyledonous plant. Ostiolar
openings are visible throughout the specimen but mature ascospores were found
only in one region. The specimen of <italic>S. lactea</italic> comprises an effused
<italic>Hypocrea</italic> on the surface of what appears to be a piece of decorticated
wood. The specimen is glued to a piece of stiff paper. The single stroma in
this collection is mostly immature with no outward evidence of ostiolar
openings or perithecia; most of it is cream or off-white in colour.
Perithecial openings are visible in one small part of the stroma; this part of
the stroma is darker in colour, more yellow, and the ostioles appear as minute
papillae. Perithecia in this region contain asci with developing ascospores,
and a few asci were found with apparently mature ascospores. It is unlikely
that the specimen identified as <italic>H. lactea</italic> is part of the original
gathering of this species simply because the genus <italic>Hypocrea</italic> dates
from 1849.</p><p>The protologues of <italic>H. citrina</italic> and <italic>H. lactea</italic> are
distinctly different, yet these taxa are now considered synonymous. According
to the protologues, <italic>H. citrina</italic> is yellow and effuse and occurs on
leaf litter, while <italic>H. lactea</italic> is white and effuse and occurs on soil
and bark. As early as 1818, Fries's concept of <italic>S. lactea</italic> began to
change as he illustrated <italic>S. lactea</italic> as a goldish white to greyish
orange effuse fungus and suggested that it was closely related to <italic>H.
citrina</italic>. Further confusing the issue, Doi's
(<xref ref-type="bibr" rid="ref9">1972</xref>) concept of <italic>H.
lactea</italic>, which is based on Japanese material, differs from Rifai &
Webster's concept of <italic>H. lactea.</italic> Doi's
(<xref ref-type="bibr" rid="ref9">1972</xref>) description and
illustration of <italic>H. lactea</italic> featuring a yellow-brown stroma with
distinct hyphal elements near the surface, likely represents an undescribed
species from Japan. The importance that Fries
(<xref ref-type="bibr" rid="ref11">1816</xref>) and Persoon
(<xref ref-type="bibr" rid="ref27">1796</xref>) ascribed to stroma
colour in the protologues of <italic>H. lactea</italic> and <italic>H. citrina</italic> must
be reexamined using current methods including molecular phylogenetics. By
designating the specimen in Fries's herbarium labeled <italic>S. lactea</italic> as
neotype, we intend to stabilize the application of the name <italic>H. lactea</italic>and provide the basis for comparing <italic>H. lactea</italic> and <italic>H.
citrina</italic>.</p></sec><sec><title>Species recognition</title><p>Species limits in <italic>Hypocrea</italic> have been historically based on
morphology. Phylogenetic data and genealogical concordance phylogenetic
species recognition (GCPSR: <xref ref-type="bibr" rid="ref32">Taylor <italic>et
al.</italic> 2000</xref>) are now being employed to define species and for
systematic studies of <italic>Hypocrea</italic> (Chaverri <italic>et al.</italic><xref ref-type="bibr" rid="ref6">2001</xref>,
<xref ref-type="bibr" rid="ref4">2003a</xref>). GCPSR utilizes the
concordance of multiple gene genealogies and the transition from concordance
among branches to incongruity among branches to establish species limits
(<xref ref-type="bibr" rid="ref32">Taylor <italic>et al.</italic> 2000</xref>).
Molecular data from RNA polymerase II subunit (<italic>rpb2</italic>), translation
elongation factor 1-alpha (<italic>tef1</italic>), and ITS1-5.8S-ITS2 (ITS) rDNA, have
been used to make crucial anamorph–teleomorph connections in
<italic>Hypocrea</italic>, to establish anamorph relationships at the sectional level
in <italic>Trichoderma</italic>, and to place genera previously described in other
pyrenomycete orders within the <italic>Hypocreales</italic>(<xref ref-type="bibr" rid="ref5">Chaverri <italic>et al.</italic>2003b</xref>, <xref ref-type="bibr" rid="ref8">Dodd <italic>et al.</italic>2002</xref>, <xref ref-type="bibr" rid="ref19">Kullnig-Gradinger <italic>et
al.</italic> 2002</xref>, <xref ref-type="bibr" rid="ref35">Zhang &
Blackwell 2002</xref>). Data from these gene regions and morphological
analyses will be used to address the following objectives: (1) the
typification and redescription of <italic>H. citrina sensu</italic> Persoon and <italic>H.
lactea sensu</italic> Fries using strict interpretations of the protologues for
typification; (2) the determination of species related to <italic>Hypocrea
citrina</italic> using GCPSR; (3) the morphological description of phylogenetic
species identified in this study; and (4) a discussion of several species
treated by Doi (<xref ref-type="bibr" rid="ref9">1972</xref>) in
<italic>Hypocrea</italic> subsect. <italic>Citrinae</italic> and subsect.
<italic>Pulvinatae</italic>.</p></sec></sec><sec sec-type="materials|methods"><title>MATERIALS AND METHODS</title><sec><title>Collections and isolates</title><p>Species concepts, illustrations, and descriptions from Doi
(<xref ref-type="bibr" rid="ref9">1972</xref>) were used in making
identifications except for <italic>H. lactea</italic> and <italic>H. fulva</italic>.
<xref ref-type="table" rid="tbl1">Table 1</xref> lists the isolates used
in this study. Frequently cited collectors are abbreviated, B.E. Overton
(B.E.O.) and G. J. Samuels (G.J.S.). 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
annotated, host and substrate information were obtained from herbarium labels,
and data were entered in <italic>specimens examined</italic> as it appears on the
labels.</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>Isolate 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><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"> DQ835433
</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"> DQ835429
</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"> DQ835430
</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
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835407
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835431
</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"> DQ835434
</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"> DQ835432
</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. Põldmaa 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"> DQ835428
</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"> DQ835427
</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.) Fr.
</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"> DQ835437
</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"> DQ835436
</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, England, U.K.
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835414
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835438
</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="center" valign="top"> DQ835442
</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"> Baarn, Zandheuvelweg, Netherlands
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835415
</td><td colspan="1" rowspan="1" align="center" valign="top"> DQ835439
</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="center" valign="top"> DQ835440
</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"> DQ835441
</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="center" valign="top"> DQ835443
</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"> DQ835444
</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="center" valign="top"> DQ835445
</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"> DQ835446
</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"> DQ835447
</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="right" valign="top"> -
</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"> Chattahoochee National Forest, 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"> DQ835449
</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"> Mt. Lake Biological Station, 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"> DQ835450
</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">Brown County, 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">DQ835448
</td><td colspan="1" rowspan="1" align="center" valign="top">AF545560
</td></tr></tbody></table></table-wrap></p></sec><sec><title>Molecular phylogenetic analysis</title><p>DNA sequence analysis of the isolates was conducted using three gene
sequences: ITS1-5.8S-ITS2 (ITS), a partial sequence of translation elongation
factor (<italic>tef1</italic>), delimited by the primer pair ef1/2, and a partial
sequence of RNA polymerase II subunit (<italic>rpb2</italic>) delimited by the primer
pair frpb2-5f/-7cr. Mycelia were lyophilized prior to DNA extraction. The
lyophilization protocol is described in Stewart <italic>et al.</italic>(<xref ref-type="bibr" rid="ref31">1999</xref>). Extraction of genomic
DNA was carried out using the phenol and chloroform extraction outlined in
Stewart <italic>et al.</italic>(<xref ref-type="bibr" rid="ref31">1999</xref>). A 50 μL polymerase
chain reaction (PCR) for ITS and <italic>tef1</italic> was performed following the
conditions outlined by Chaverri <italic>et al.</italic>(<xref ref-type="bibr" rid="ref6">2001</xref>) using the following
primer pairs: for ITS, the primers utilized were ITS1
(5'-TCCGTAGGTGAACCTGCGG-3') and ITS4 (5'-TCCTCCGCTTATTGATATGC-3')
(<xref ref-type="bibr" rid="ref34">White <italic>et al.</italic> 1990</xref>);
for <italic>tef1</italic>, the primers were ef-1 (5'-ATGGGTAAGGA(A/G)GACAAGAC-3') and
ef-2 (5'-GGA(G/A)GTACCAGT(G/C)ATCATGTT-3')
(<xref ref-type="bibr" rid="ref24">O'Donnell <italic>et al.</italic>1998</xref>). A 50 μL polymerase chain reaction (PCR) following the
conditions outlined by Chaverri <italic>et al.</italic>(<xref ref-type="bibr" rid="ref5">2003b</xref>) was utilized for
generating a partial <italic>rpb2</italic> product. The following two fungus-specific
primers were employed: frpb2-5f
(5'-GA(T/C)GA(T/C)(A/C)G(A/T)GATCA(T/C)TT(T/C)GG -3') and
frpb2-7cr(5'-CCCAT(A/G)GCTTG(T/C)TT(A/G)CCCAT-3')
(<xref ref-type="bibr" rid="ref20">Liu <italic>et al.</italic> 1999</xref>).
Two percent dimethyl sulfoxide (DMSO) from AMRESCO® was added to each 50
μL PCR reaction. A PCR purification was carried out using a QIAquick®
PCR purification kit. The ABI Prism Big Dye Terminator Cycle Sequencing Ready
Reaction Kit with Amplitaq DNA Polymerase FS (PE Applied Biosystems) was
utilized for sequencing cleaned PCR products for each primer direction.
Performa® DTR gel filtration cartridges from Edge BioSystems were used for
cleaning cycle sequencing products. Sequencing was either conducted on an ABI
377 located in the Plant Pathology Department at The Pennsylvania State
University or sent to the Nucleic Acid Facility (Life Science Consortium, The
Pennsylvania State University). Sequence data were trimmed at the 5' and 3'
ends. The ITS region was sequenced for all isolates and used to screen for
replicate isolates. Replicate isolates were removed from subsequent analyses
if there were no obvious morphological or locality differences noted from the
herbarium specimens. 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. 4.0b8 (Sinauer Associates,
Sunderland, MA). Alignments were manually adjusted in PAUP*. Overton <italic>et
al.</italic> (<xref ref-type="bibr" rid="ref25">1999</xref>) determined
<italic>H. sulphurea</italic> to be a phylogenetically appropriate outgroup taxon for
<italic>H. citrina</italic> and allies and this species was used as outgroup in all
data sets. Maximum parsimony (MP) analyses were completed using the heuristic
search option with starting trees obtained via random addition sequence (10
replicates) and TBR branch swapping. Gaps (insertions/deletions) were treated
as missing data. Bootstrap analysis was performed in 500 replicates with
random sequence addition (10 replicates). The sequences and alignments were
deposited in GenBank (<xref ref-type="table" rid="tbl1">Table
1</xref>).</p></sec><sec><title>Determination of optimal growth temperature</title><p>A growth trial using multiple isolates from each phylogenetic species
established in the species tree (<xref rid="fig1" ref-type="fig">Fig.
1</xref>) was carried out using 20 mL of three media: potato-dextrose
agar (PDA, Difco), cornmeal-dextrose agar (CMD, Difco), and SNA with filter
paper (<xref ref-type="bibr" rid="ref23">Nirenberg 1976</xref>).
Inoculum was prepared on CMD. A 1-cm-diam disk from the actively growing
inoculum colony was placed near the edge of the plate. One plate of each
medium for each culture was incubated at 20, 25, 30, and 35 °C in total
darkness. The radius of the colony was measured from the edge of the inoculum
at 24-h intervals for 4 d. The experiment was replicated three times during
subsequent weeks. Optimal growth temperatures are summarized in the species
descriptions.</p></sec><sec><title>Morphological descriptions</title><p>Anamorph and teleomorph characteristics were measured from isolates and
specimens representative of each phylogenetic species distinguished in
<xref rid="fig1" ref-type="fig">Fig. 1</xref>. Cultures of
<italic>Hypocrea</italic> were grown on PDA, CMD and SNA at 20 °C, with 12 h
fluorescent light and 12 h darkness to observe and measure microscopic
characters of the anamorph. Observations were made at ∼7–10 d.
Observations were discontinued for cultures that did not produce conidiophores
after 10 d on each respective medium. Anamorph characters measured were
phialide length, width of the phialides at the middle, and conidium length and
width. Phialide arrangement, number of phialides in a whorl and colony
characteristics were recorded. Herbarium specimens were rehydrated in 3 % KOH
for sectioning. Rehydrated stromata were supported by Tissue-Tek O.C.T.
Compound 4583 (Miles Inc., Elkhart, Indiana) and sectioned at a thickness of
approximately 15 μm with a freezing microtome. Teleomorph characters
measured were ascus length, ascus width, ascospore proximal length and width,
ascospore distal length and width, perithecial length and width, and stroma
length and width. Stroma colour, including colour reactions of stroma tissue
in response to 3 % KOH and lactic acid, were also recorded. Anatomical
characters of the stroma were described and photographed. Measurements of
continuous characters were made using the image-capturing software Scion Image
beta 4.0.2 (Scion Corporation, Frederick, Maryland). Descriptive statistics of
micromorphological characters were made based on 30 measurements per specimen,
except where noted, with confidence intervals (a = 0.05), and minimum and
maximum values reported. Colour terminology was obtained from Kornerup &
Wanscher (<xref ref-type="bibr" rid="ref18">1981</xref>). Anamorph and
teleomorph characters are included in the species descriptions. Important
morphological characters used in species recognition are discussed in the
comments section immediately following each species description.</p></sec></sec><sec><title>RESULTS</title><p>The trees generated from ITS are slightly different from those obtained
from <italic>tef1</italic> and <italic>rpb2</italic> (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>). In the ITS gene tree, the
proposed neotype of <italic>H. citrina</italic> from Europe
(<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>)
differed from North American isolates of <italic>H. citrina</italic> in a single
nucleotide. The <italic>H. citrina</italic> isolate G.J.S. 95-183 is identical to
other isolates from North America in the ITS tree. The ITS result differed
from the <italic>tef1</italic> gene tree in which the European
<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> was
identical with the North American isolate B.E.O. 99-29
(<xref rid="fig2" ref-type="fig">Fig. 2</xref>) and from the
<italic>tef1</italic> and <italic>rpb2</italic> gene trees, in which isolate G.J.S. 95-183 had
a longer branch length than other isolates of <italic>H. citrina</italic> (Figs
<xref rid="fig2" ref-type="fig">2</xref>,
<xref rid="fig3" ref-type="fig">3</xref>). Parsimony analysis of the
ITS sequences showed a single nucleotide polymorphism separating isolates of
<italic>H. pulvinata</italic> from <italic>H. americana</italic> supported by a bootstrap
score of 63 % (<xref rid="fig1" ref-type="fig">Fig. 1</xref>). The
<italic>tef1</italic> and <italic>rpb2</italic> gene trees were more pronounced in placing
<italic>H. citrina</italic> in a highly supported monophyletic group with <italic>H.
pulvinata, H. protopulvinata</italic>, and <italic>H. americana</italic> with bootstrap
scores of 99 % for both trees.</p><p>All three data sets are concordant with similar homoplasy indices. The
heuristic search of the most parsimonious tree for the ITS dataset yielded two
trees with 48 steps. The minimal possible tree length is 41
(<xref rid="fig1" ref-type="fig">Fig. 1</xref>). From a total of 523
characters, 484 characters are constant, 11 variable characters are
parsimony-uninformative and 28 characters are parsimony-informative. The
heuristic search of the most parsimonious trees for the <italic>tef1</italic> dataset
yielded six trees with 476 steps with the minimum possible tree length of 393
(<xref rid="fig2" ref-type="fig">Fig. 2</xref>). From 695 total
characters, 392 characters are constant, 61 variable characters are
parsimony-uninformative, and 242 characters are parsimony-informative. The
heuristic search of the most parsimonious trees for the <italic>rpb2</italic> dataset
yielded two trees with 172 steps with the minimum possible tree length of 150
(<xref rid="fig3" ref-type="fig">Fig. 3</xref>). From 969 total
characters, 833 characters are constant, 56 variable characters are
parsimony-uninformative, and 80 characters are parsimony-informative.</p><p>The combined phylogenetic analysis using ITS, partial sequences of
<italic>tef1</italic> and <italic>rpb2</italic> distinguished <italic>H. americana</italic> from
<italic>H. pulvinata</italic> with a bootstrap score of 100 %
(<xref rid="fig4" ref-type="fig">Fig. 4</xref>). European and North
American isolates of <italic>H. pulvinata</italic> formed two distinct subclades, each
supported by a bootstrap score of 100 %. <italic>Hypocrea pulvinata</italic> and
<italic>H. americana</italic> formed a monophyletic group supported by a bootstrap
score of 87 % (I, II). <italic>Hypocrea americana, H. pulvinata</italic>, and <italic>H.
protopulvinata</italic> (III) formed a strongly supported monophyletic group with
a bootstrap score of 100 % (Clade A). Isolates of <italic>H. citrina</italic>(including the ex-neotype (IV)) formed a strongly supported monophyletic group
with a bootstrap score of 100 % (Clade B). <italic>Hypocrea microcitrina</italic> (VI)
and <italic>H. pseudostraminea</italic> (V) are situated in an unresolved sister clade
to <italic>H. citrina</italic> supported by a bootstrap score of 100 % (Clade C).
Isolate B.E.O. 99-36 is divergent from other isolates of <italic>H.
pseudostraminea</italic>; nevertheless, all isolates of <italic>H.
pseudostraminea</italic> formed a strongly supported monophyletic group (V) with a
bootstrap score of 100 %. Isolates of <italic>H. microcitrina</italic> (VI) formed a
strongly supported clade with a bootstrap score of 100 %. The heuristic search
of the most parsimonious trees yielded one tree with 678 steps with the
minimum possible tree length of 575 (<xref rid="fig4" ref-type="fig">Fig.
4</xref>). From 2189 total characters, 1710 characters are constant, 133
variable characters are parsimony-uninformative, and 346 characters are
parsimony-informative.</p><p>Overall, ITS sequences are less variable than the <italic>tef1</italic> and
<italic>rpb2</italic> sequences. The <italic>tef1</italic> and <italic>rpb2</italic> regions
distinguished between North American and European isolates of <italic>H.
pulvinata</italic>, whereas ITS did not. None of the gene regions consistently
resolved North American and European isolates of <italic>H. citrina. Hypocrea
citrina</italic> isolate G.J.S. 95-183 grouped with North American and European
isolates of <italic>H. citrina</italic> in the ITS tree, but had a significantly
longer branch in the <italic>tef1</italic> and <italic>rpb2</italic> trees. This represents a
point of discordance that establishes a phylogenetic species boundary for
<italic>H. citrina</italic>. The highest bootstrap scores for isolates of <italic>H.
citrina</italic> were obtained in all three datasets at the base of Clade B.</p></sec><sec><title>DISCUSSION</title><sec><title>Phylogenetic analysis and establishment of major clades</title><p>Despite some variation between the ITSl, <italic>tef1</italic>, and <italic>rpb2</italic>gene trees, it can be concluded that <italic>H. citrina</italic> is phylogenetically
distinct from <italic>H. pulvinata</italic>. In the combined analysis, Clade A,
consisting of <italic>H. pulvinata, H. americana,</italic> and <italic>H.
protopulvinata,</italic> is monophyletic; Clade B consists of all <italic>H.
citrina</italic> isolates; and Clade C consists of <italic>H. pseudostraminea</italic> and
<italic>H. microcitrina</italic> (<xref rid="fig4" ref-type="fig">Fig.
4</xref>). The <italic>H. microcitrina</italic> and <italic>H. pseudostraminea</italic>subclades are separated by long branches
(<xref rid="fig4" ref-type="fig">Fig. 4</xref>). It is possible, given
the large branch length differences, that long-branch attraction may be
artificially inflating the bootstrap score uniting these taxa. Future
phylogenetic studies that should include <italic>H. protocitrina, H.
albocitrina</italic>, and <italic>H. lactea sensu</italic> Doi may help to more clearly
elucidate the relationship of <italic>H. pseudostraminea</italic> and <italic>H.
microcitrina</italic> to <italic>H. citrina</italic>.</p><p>Sequences from <italic>H. pulvinata</italic> and <italic>H. americana</italic> produced
highly concordant gene trees, consistently generating the same subgroupings.
ITS data from 3 out of 4 isolates of <italic>H. citrina</italic> from Europe differed
from North American isolates in a single nucleotide. The ex-neotype culture of
<italic>H. citrina</italic> (<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>) was identical with B.E.O. 99-29 in the <italic>tef1</italic> gene
tree, but they differed in the <italic>rpb2</italic> tree. Furthermore, <italic>H.
citrina</italic> isolate G.J.S. 95-183, which is invariant in the ITS tree from
other North American isolates, had significant branch length differences in
the <italic>tef1</italic> and <italic>rpb2</italic> gene trees from all other <italic>H.
citrina</italic> isolates. The differences between isolates of <italic>H. citrina</italic>can best be explained by sexual recombination. Therefore, the species boundary
for <italic>H. citrina</italic> was determined to be at the point of discordance
separating clade B in the gene trees (<xref rid="fig4" ref-type="fig">Fig.
4</xref>). Isolates of <italic>H. pulvinata</italic> show large numbers of
nucleotide changes between North American and European isolates in the
<italic>rpb2</italic> and <italic>tef1</italic> data sets. The differences between North
American and European isolates in the <italic>tef1</italic> and <italic>rpb2</italic> may
represent cryptic species or simply variations in population structure. North
American and European isolates of <italic>H. pulvinata</italic> are morphologically
indistinguishable, and the ITS sequences are identical. Additional sequence
analysis and morphological examination of European isolates will be required
to determine whether European isolates represent a distinct species. In the
ITS gene tree for <italic>H. americana</italic>, a single nucleotide change separates
isolates of <italic>H. americana</italic> from <italic>H. pulvinata</italic>, while there are
larger differences in the <italic>tef1</italic> and <italic>rpb2</italic> gene trees between
these species.</p><p><fig position="float" id="fig1"><label>Fig. 1.</label><caption><p>Parsimony analysis of ITS1-5.8S-ITS2. One phylogram of two most
parsimonious trees; 48 steps; consistency index: 0.854; retention index;
0.989; homoplasy index: 0.146; numerical values of branch lengths are given
above and bootstrap values (500 replicates with 10 random addition
replications) are indicated below branches. Outgroup taxon: <italic>H.
sulphurea</italic>.</p></caption><graphic xlink:href="1fig1"></graphic></fig></p><p>The optimal temperature for growth was determined for each phylogenetic
species. Optimal temperature for growth is reported only when multiple strains
with branch length differences within a phylogenetic species could be sampled
in order to determine the intra-species variation. Growth temperature optima
could not adequately distinguish between any of the phylogenetic species nor
were they variable within a phylogenetic species in which strain differences
had been established based on branch length differences.</p></sec><sec><title>Phylogenetics and classification</title><p>The monophyly of many of the infrageneric groupings proposed by Doi
(<xref ref-type="bibr" rid="ref9">1972</xref>) have yet to be tested by
DNA sequence analysis. In this study, it was found that subsections
<italic>Citrinae</italic> and <italic>Pulvinatae</italic> as defined by Doi are not
monophyletic assemblages of species. <italic>Hypocrea microcitrina</italic> and <italic>H.
megalocitrina</italic> (<xref ref-type="bibr" rid="ref5">Chaverri <italic>et
al.</italic> 2003b</xref>) are not situated in the <italic>H. citrina</italic> clade.
It is more difficult to determine the phylogenetic boundaries of Subsection
<italic>Pulvinatae</italic> because Doi broke it up into several series from which
isolates could not be sequenced.</p><p><fig position="float" id="fig2"><label>Fig. 2.</label><caption><p>Parsimony analysis of partial sequences of <italic>tef1</italic>. One phylogram of
six most parsimonious trees; 476 steps; consistency index: 0.826; retention
index: 0.769; homoplasy index: 0.174. Rest as in
<xref rid="fig1" ref-type="fig">Fig. 1</xref>.</p></caption><graphic xlink:href="1fig2"></graphic></fig></p><p><italic>Hypocrea microcitrina, H. pseudostraminea</italic> and <italic>H. lactea
sensu</italic> Doi have similar morphological characters in the form of hyphal
protrusions and <italic>textura intricata</italic> near the stroma surface. It is
difficult to distinguish between hyphal protuberances and <italic>textura
intricata</italic>. It is unclear at this time if the hyphal protrusions near the
surface of the stromata and the <italic>textura intricata</italic> found in <italic>H.
pseudostraminea</italic> and <italic>H. microcitrina</italic> are homologous characters,
but both of these taxa have similar irregular verticillium-like anamorphs. Doi
(<xref ref-type="bibr" rid="ref9">1972</xref>) classified <italic>H.
pseudostraminea</italic> in subsect. <italic>Pulvinatae</italic>, series
<italic>Splendentes</italic>, and <italic>H. microcitrina</italic> in subsect.
<italic>Citrinae</italic>. The results generated here suggest that the subgroupings
proposed by Doi (<xref ref-type="bibr" rid="ref9">1972</xref>) are
artificial (<xref ref-type="table" rid="tbl2">Table 2</xref>).</p><p><table-wrap position="float" id="tbl2"><label>Table 2.</label><caption><p>Phylogenetic clades and morphological characters.</p></caption><table frame="hsides" rules="groups"><thead><tr><th colspan="1" rowspan="1" align="left" valign="top"><bold>Species</bold></th><th colspan="1" rowspan="1" align="center" valign="top"><bold>Clade</bold></th><th colspan="1" rowspan="1" align="left" valign="top"><bold>Ascospore ornamentation</bold></th><th colspan="1" rowspan="1" align="left" valign="top"><bold>Stroma anatomy (textura)</bold></th><th colspan="1" rowspan="1" align="left" valign="top"><bold>Hyphal protuberances above stroma surface</bold></th><th colspan="1" rowspan="1" align="left" valign="top"><bold>Anamorph and conidial shape</bold></th><th colspan="1" rowspan="1" align="left" valign="top"><bold>Infrageneric classification (</bold><xref ref-type="bibr" rid="ref9"><bold>Doi
1972</bold></xref><bold>) in <italic>Hypocrea</italic> subgenus <italic>Hypocrea</italic>Section <italic>Homalocrea</italic></bold></th></tr></thead><tbody><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>H. pulvinata</italic></td><td colspan="1" rowspan="1" align="center" valign="top"> A
</td><td colspan="1" rowspan="1" align="left" valign="top"> smooth to minutely spinulose
</td><td colspan="1" rowspan="1" align="left" valign="top"><italic>t. globulosa</italic> to <italic>t. angularis</italic> with vertically parallel <italic>t.
intricata</italic> (compact)
</td><td colspan="1" rowspan="1" align="left" valign="top"> warted hyphae
</td><td colspan="1" rowspan="1" align="left" valign="top"> acremonium-like subglobose—subellipsoidal
</td><td colspan="1" rowspan="1" align="left" valign="top"> Subsection <italic>Pulvinatae</italic> series <italic>Pulvinatae</italic></td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>H. americana</italic></td><td colspan="1" rowspan="1" align="center" valign="top"> A
</td><td colspan="1" rowspan="1" align="left" valign="top"> smooth
</td><td colspan="1" rowspan="1" align="left" valign="top"><italic>t. globulosa</italic> to <italic>t. angularis</italic> with vertically parallel <italic>t.
intricata</italic> (compact)
</td><td colspan="1" rowspan="1" align="left" valign="top"> warted hyphae
</td><td colspan="1" rowspan="1" align="left" valign="top"> acremonium-like subglobose—subellipsoidal
</td><td colspan="1" rowspan="1" align="left" valign="top"> Subsection <italic>Pulvinatae</italic> series <italic>Pulvinatae</italic></td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>H. protopulvinata</italic></td><td colspan="1" rowspan="1" align="center" valign="top"> A
</td><td colspan="1" rowspan="1" align="left" valign="top"> smooth
</td><td colspan="1" rowspan="1" align="left" valign="top"><italic>t. globulosa</italic> with vertically parallel <italic>t. intricata</italic> (not
compact)
</td><td colspan="1" rowspan="1" align="left" valign="top"> roughened hyphae
</td><td colspan="1" rowspan="1" align="left" valign="top"> acremonium-like subglobose—subellipsoidal
</td><td colspan="1" rowspan="1" align="left" valign="top"> Subsection <italic>Pulvinatae</italic> series <italic>Pulvinatae</italic></td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>H. platypulvinata</italic></td><td colspan="1" rowspan="1" align="center" valign="top"> likely clade A
</td><td colspan="1" rowspan="1" align="left" valign="top"> smooth to minutely spinulose
</td><td colspan="1" rowspan="1" align="left" valign="top"><italic>t. globulosa</italic> to <italic>t. angularis</italic> with vertically parallel <italic>t.
intricata</italic> (compact)
</td><td colspan="1" rowspan="1" align="left" valign="top"> warted hyphae
</td><td colspan="1" rowspan="1" align="left" valign="top"> acremonium-like subglobose—cylindrical
</td><td colspan="1" rowspan="1" align="left" valign="top"> Subsection <italic>Pulvinatae</italic> series <italic>Pulvinatae</italic></td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>H. citrina</italic></td><td colspan="1" rowspan="1" align="center" valign="top"> B
</td><td colspan="1" rowspan="1" align="left" valign="top"> minutely spinulose
</td><td colspan="1" rowspan="1" align="left" valign="top"><italic>t. globulosa</italic> to <italic>t. angularis</italic> with some vertically parallel
<italic>t. intricata</italic> (compact)
</td><td colspan="1" rowspan="1" align="left" valign="top"> none
</td><td colspan="1" rowspan="1" align="left" valign="top"> verticillium-like subglobose—subellipsoidal
</td><td colspan="1" rowspan="1" align="left" valign="top"> Subsection <italic>Citrinae</italic></td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>H. pseudostraminea</italic></td><td colspan="1" rowspan="1" align="center" valign="top"> C
</td><td colspan="1" rowspan="1" align="left" valign="top"> minutely spinulose
</td><td colspan="1" rowspan="1" align="left" valign="top"> vertically parallel <italic>t. intricata</italic> (compact) with thin layer of <italic>t.
globulosa</italic> to <italic>t. angularis</italic> near stroma surface
</td><td colspan="1" rowspan="1" align="left" valign="top"> roughened hyphae
</td><td colspan="1" rowspan="1" align="left" valign="top"> verticillium-like subellipsoidal—elongate ellipsoidal
</td><td colspan="1" rowspan="1" align="left" valign="top"> Subsection <italic>Pulvinatae</italic> Series <italic>Splendentes</italic></td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>H. microcitrina</italic></td><td colspan="1" rowspan="1" align="center" valign="top"> C
</td><td colspan="1" rowspan="1" align="left" valign="top"> minutely spinulose
</td><td colspan="1" rowspan="1" align="left" valign="top"> vertically parallel <italic>t. intricata</italic> (compact to loosely woven)
</td><td colspan="1" rowspan="1" align="left" valign="top"><italic>t.
intricata</italic><xref ref-type="table-fn" rid="tblfn2">2</xref></td><td colspan="1" rowspan="1" align="left" valign="top"> verticillium-like subellipsoidal—elongate ellipsoidal
</td><td colspan="1" rowspan="1" align="left" valign="top"> Subsection <italic>Citrinae</italic></td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>H. lactea sensu</italic> Doi
</td><td colspan="1" rowspan="1" align="center" valign="top"> likely clade C
</td><td colspan="1" rowspan="1" align="left" valign="top"> spinulose
</td><td colspan="1" rowspan="1" align="left" valign="top"> vertically parallel <italic>t. intricata</italic> (compact to loosely woven)
</td><td colspan="1" rowspan="1" align="left" valign="top"><italic>t. intricata</italic> roughened near stroma surface
</td><td colspan="1" rowspan="1" align="right" valign="top"> —
</td><td colspan="1" rowspan="1" align="left" valign="top"> Subsection <italic>Citrinae</italic></td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>H. megalocitrina</italic></td><td colspan="1" rowspan="1" align="center" valign="top"><italic>Psychrophila</italic><xref ref-type="table-fn" rid="tblfn1">1</xref>clade
</td><td colspan="1" rowspan="1" align="left" valign="top"> nodulose-warted
</td><td colspan="1" rowspan="1" align="left" valign="top"> vertically parallel <italic>t. intricata</italic> (compact to loosely woven)
</td><td colspan="1" rowspan="1" align="left" valign="top"><italic>t.
intricata</italic><xref ref-type="table-fn" rid="tblfn2">2</xref></td><td colspan="1" rowspan="1" align="left" valign="top"> verticillium-like subglobose—ellipsoidal, sometimes elongate ellipsoidal
</td><td colspan="1" rowspan="1" align="left" valign="top"> Subsection <italic>Citrinae</italic></td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>H. psychrophila</italic></td><td colspan="1" rowspan="1" align="center" valign="top"><italic>psychrophila</italic><xref ref-type="table-fn" rid="tblfn1">1</xref>clade
</td><td colspan="1" rowspan="1" align="left" valign="top"> nodulose-warted
</td><td colspan="1" rowspan="1" align="left" valign="top"> thick-walled cells (compact), <italic>t. angularis</italic></td><td colspan="1" rowspan="1" align="left" valign="top"> none
</td><td colspan="1" rowspan="1" align="left" valign="top"> gliocladium-like ovoid to subellipsoidal
</td><td colspan="1" rowspan="1" align="left" valign="top"> Not included by Doi
</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>H. fulva</italic></td><td colspan="1" rowspan="1" align="center" valign="top"> likely <italic>psychrophila</italic> clade
</td><td colspan="1" rowspan="1" align="left" valign="top"> nodulose-warted
</td><td colspan="1" rowspan="1" align="left" valign="top"> vertically parallel <italic>t. intricata</italic> (compact)
</td><td colspan="1" rowspan="1" align="left" valign="top"> none (thin layer of compact cells at surface)
</td><td colspan="1" rowspan="1" align="right" valign="top"> —
</td><td colspan="1" rowspan="1" align="left" valign="top"> Subsection <italic>Citrinae</italic></td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>H. mikurajimensis</italic></td><td colspan="1" rowspan="1" align="center" valign="top"> likely <italic>psychrophila</italic> clade
</td><td colspan="1" rowspan="1" align="left" valign="top"> nodulose-warted
</td><td colspan="1" rowspan="1" align="left" valign="top"> thick-walled cells (compact)
</td><td colspan="1" rowspan="1" align="left" valign="top"> none
</td><td colspan="1" rowspan="1" align="left" valign="top"> verticillium-like ovoid to subellipsoidal
</td><td colspan="1" rowspan="1" align="left" valign="top"> Subsection <italic>Pulvinatae</italic> series <italic>Splendentes</italic></td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"><italic>H. aurantiistroma</italic></td><td colspan="1" rowspan="1" align="center" valign="top">likely <italic>psychrophila</italic> clade
</td><td colspan="1" rowspan="1" align="left" valign="top">minutely nodulose
</td><td colspan="1" rowspan="1" align="left" valign="top">vertically parallel <italic>t. intricata</italic> (loose)
</td><td colspan="1" rowspan="1" align="left" valign="top"><italic>t.
intricata</italic><xref ref-type="table-fn" rid="tblfn2">2</xref></td><td colspan="1" rowspan="1" align="right" valign="top">—
</td><td colspan="1" rowspan="1" align="right" valign="top">—
</td></tr></tbody></table><table-wrap-foot><fn id="tblfn1"><label>1</label><p>The <italic>psychrophila</italic> clade is supported by a bootstrap score of 93 %
based on combined parsimony analysis of <italic>rpb2</italic> and <italic>tef1</italic>(<xref ref-type="bibr" rid="ref5">Chaverri <italic>et al.</italic>2003b</xref>, fig. 7).</p></fn><fn id="tblfn2"><label>2</label><p>It is difficult to distinguish between hyphal protuberances and <italic>t.
intricata</italic> near the stroma surface because <italic>t. intricata</italic> does not
form a well-defined margin at the stroma surface resulting in hyphal
protuberances of varying lengths.</p></fn></table-wrap-foot></table-wrap></p><p>At this time, it is unclear how <italic>H. albocitrina</italic> and <italic>H.
protocitrina</italic> are related to <italic>H. citrina</italic>, as cultures are not
available for sequencing. The overall macromorphology of <italic>H.
albocitrina</italic> and the stroma tissue of <italic>H. protocitrina</italic> consisting
of <italic>textura intricata</italic> suggest that these taxa are more similar to
<italic>H. microcitrina</italic> and <italic>H. pseudostraminea. Hypocrea
megalocitrina</italic> has been sequenced and is not situated in the same clade as
<italic>H. citrina</italic> (<xref ref-type="bibr" rid="ref5">Chaverri <italic>et
al.</italic> 2003b</xref>). Molecular data generated by Chaverri <italic>et
al.</italic> (<xref ref-type="bibr" rid="ref5">2003b</xref>) place <italic>H.
megalocitrina</italic> near a cold-loving species described by Müller <italic>et
al.</italic> (<xref ref-type="bibr" rid="ref21">1972</xref>) as <italic>H.
psychrophila</italic>. Doi (<xref ref-type="bibr" rid="ref10">2001</xref>)
described a species from Japan, <italic>H. mikurajimensis</italic> Yoshim. Doi that
has characters similar to those of <italic>H. fulva, H. megalocitrina, H.
aurantiistroma</italic>, and <italic>H. psychrophila</italic>. The strongly warted
ascospores, stroma anatomy, and yellow pulvinate stromata of <italic>H.
mikurajimensis</italic> are similar to the characteristics described by
Müller <italic>et al.</italic>(<xref ref-type="bibr" rid="ref21">1972</xref>) for <italic>H.
psychrophila,</italic> which has a white gliocladium-like anamorph. The nodulose
ascospores of <italic>H. mikurajimensis</italic>, longitudinal elongate ostiolar
canals, and regularly verticillate conidiophores are similar to characters
found in <italic>H. megalocitrina, H. aurantiistroma</italic>, and <italic>H. fulva</italic>.
The warted, thick-walled ascospores and long ostiolar canals of <italic>Hypocrea
psychrophila</italic> and <italic>H. megalocitrina</italic> are the morphological
characters uniting these taxa.</p><p><fig position="float" id="fig3"><label>Fig. 3.</label><caption><p>Parsimony analysis of partial sequences of <italic>rpb2</italic>. One phylogram of
two most parsimonious trees; 172 steps; consistency index: 0.872; retention
index: 0.914; homoplasy index: 0.128. Rest as in
<xref rid="fig1" ref-type="fig">Fig. 1</xref>.</p></caption><graphic xlink:href="1fig3"></graphic></fig></p><p>Doi (<xref ref-type="bibr" rid="ref10">2001</xref>) classified
<italic>H. mikurajimensis</italic> in the <italic>H. patella</italic> group of subsection
<italic>Pulvinatae</italic>, series <italic>Splendentes,</italic> which has been shown to be
paraphyletic (<xref ref-type="bibr" rid="ref8">Dodd <italic>et al.</italic>2002</xref>). The exact phylogenetic position of this species in relation
to all other taxa included by Doi
(<xref ref-type="bibr" rid="ref9">1972</xref>) in Series
<italic>Splendentes</italic> cannot be determined at this time. However, Chaverri
<italic>et al.</italic> (<xref ref-type="bibr" rid="ref5">2003b</xref>) have
shown that <italic>H. pezizoides</italic> Berk. & Broome, also series
<italic>Splendentes</italic>, is situated in the <italic>H. rufa</italic> clade. These authors
also showed that <italic>H. megalocitrina</italic> and <italic>H. psychrophila</italic> are
phylogenetically related. Based on characters of the teleomorph and anamorph
mentioned above, it is possible that <italic>H. megalocitrina, H. fulva, H.
mikurajimensis, H. aurantiistroma</italic>, and <italic>H. psychrophila</italic> are
phylogenetically related, but more taxa have to be sequenced to establish the
limits of the “<italic>psychrophila</italic>” clade.</p><p><fig position="float" id="fig4"><label>Fig. 4.</label><caption><p>Combined parsimony analysis of ITS, <italic>tef</italic>1, <italic>rpb</italic>2. Phylogram
of single most parsimonious tree; 678 steps; consistency index: 0.848;
retention index: 0.915; homoplasy index: 0.152. Rest as in
<xref rid="fig1" ref-type="fig">Fig. 1</xref>.</p></caption><graphic xlink:href="1fig4"></graphic></fig></p></sec><sec><title>Typification and nomenclatural conclusions</title><p>With the selection of a specimen that agrees with the protologue of <italic>S.
citrina</italic> (Gams 4031, <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>) as <bold>neotype</bold> (herein designated), the concept of the
species can be firmly attached to a litter-inhabiting fungus. It is clear from
the molecular and morphological data that <italic>H. pulvinata</italic> (= <italic>H.
citrina</italic> var. <italic>citrina sensu</italic> Canham) and <italic>H. citrina</italic>, as
neotypified here, are not synonyms and that Fries
(<xref ref-type="bibr" rid="ref13">1823</xref>) and later Canham
(<xref ref-type="bibr" rid="ref3">1969</xref>) made a taxonomic error
in expanding the species concept of <italic>S. citrina</italic> to include the
polyporicolous element. <italic>Hypocrea</italic> <italic>citrina</italic> var.
<italic>americana</italic> is elevated to species status, <italic>H. americana</italic>, based
on morphological characters and a well-delimited phylogenetic species
boundary.</p><p>Molecular data were compared from many different specimens of <italic>H.
citrina</italic> collected in different geographic locations with variations in
stroma colour and substrate. It was determined that the protologue was too
narrowly defined for <italic>H. citrina</italic>. Specimens BPI 1107145, G.J.S. 89-145
and <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>(<xref rid="fig5" ref-type="fig">Fig. 5C</xref>), which are white to
light yellow and occur on soil, were found to be phylogenetically the same as
the neotype of <italic>H. citrina</italic>, which is bright yellow and occurs on leaf
litter (<xref rid="fig6" ref-type="fig">Fig. 6</xref>). A lemon-yellow
specimen, B.E.O. 99-29, on the moss <italic>Drepanocladus uncinatus</italic> (Hedw.)
Warnst., was also found to be the same as the neotype based on molecular
results. These findings show that stroma colour and substratum are variable
characters, a finding not suggested by the original protologue of <italic>S.
citrina. Hypocrea citrina</italic> was never observed growing on polypores in this
study.</p><p>The only two available historical specimens in Fries's herbarium (UPS) for
<italic>H. lactea</italic> are not consistent with the protologue in that they are not
white, as emphasized in the original protologue of <italic>H. lactea</italic>.
Recommendations 9A.2 and 9A.3 of the ICBN clearly indicate that historical
specimens should not automatically be taken as type material if they are
discordant with the protologue. However, Fries's species concept of <italic>H.
lactea</italic> changed from the white form described in his protologue
(<xref ref-type="bibr" rid="ref11">1816</xref>) to include a whitish
gold to greyish orange form that Fries illustrated
(<xref ref-type="bibr" rid="ref12">1818</xref>) with a colour
lithograph two years after the original description and sanctioned
(<xref ref-type="bibr" rid="ref13">Fries 1823</xref>). The illustrated
specimen has almost exactly the same shape and colour as the specimen of
<italic>S. lactea</italic> considered here as the <bold>lectotype</bold> specimen
(<xref rid="fig7" ref-type="fig">Fig. 7</xref>). Taxonomically, both
specimens of <italic>H. lactea</italic> in Fries's herbarium can be identified as the
yellow form of <italic>H. citrina</italic> based on morphological characters.
Molecular data have shown that <italic>H. citrina</italic> could in fact be white and
could occur on soil just as Fries described in the protologue for <italic>H.
lactea</italic>. The neotype of <italic>H. citrina</italic> is consistent in
characteristics defined by Fries for <italic>H. lactea</italic> and is consistent with
Rifai & Webster's (<xref ref-type="bibr" rid="ref28">1966</xref>)
conclusion that <italic>H. citrina</italic> and <italic>H. lactea</italic> are synonymous.</p><p>Rifai & Webster's selection of the name <italic>H. lactea</italic> over that of
<italic>H. citrina</italic> is not consistent with the present Articles 11.4 and 15.4
of the ICBN, which govern the priority when two sanctioned names compete,
giving priority to the older name. Consequently, terrestrial collections
lodged in herbaria after 1966 were generally labeled <italic>H. lactea</italic>.
Specimens previously identified under the name <italic>H. lactea</italic> should
therefore be reidentified as <italic>H. citrina</italic>, the older name, which has
priority over <italic>H. lactea</italic>.</p><p>Some confusion exists around the selection of type material associated with
the name <italic>H. pulvinata</italic>. When Fuckel
(<xref ref-type="bibr" rid="ref15">1870</xref>) described <italic>H.
pulvinata</italic>, he did not cite a specific specimen, but indicated
<italic>Polyporus sulphureus</italic> as the substratum. Rifai & Webster
(<xref ref-type="bibr" rid="ref28">1966</xref>) indicated that the
substratum for Fuckel<italic>, Fungi Rhenani</italic> 2467, preserved at K was
<italic>“Polyporus resinosus”</italic>, which recently was identified as
<italic>Laetiporus sulphureus</italic>. This specimen is over-mature, and no asci
remain. Two specimens of <italic>F. Rhenani</italic> 2467 on <italic>L. sulphureus</italic> in
G are also overmature. A specimen at FH, #876, was part of Fuckel's herbarium;
the collecting data match those given in the protologue and the substratum
also is identified as being <italic>Polyporus</italic> (= <italic>Laetiporus</italic>)
<italic>sulphureus</italic>. Moreover, this collection is in good condition and is
designated here as the <bold>lectotype</bold> for <italic>H. pulvinata</italic>.
Measurements obtained from all historical specimens included in this study
were consistent with measurements from more recent collections of <italic>H.
pulvinata</italic> used in the molecular analysis. The species concept presented
here for the polyporicolous form and the application of the name <italic>H.
pulvinata</italic> is consistent with Weese
(<xref ref-type="bibr" rid="ref33">1927</xref>), Rifai & Webster
(<xref ref-type="bibr" rid="ref28">1966</xref>), and Doi
(<xref ref-type="bibr" rid="ref9">1972</xref>).</p></sec></sec><sec><title>TAXONOMY</title><p><list list-type="order"><list-item><p><bold><italic>Hypocrea citrina</italic></bold> (Pers.: Fr.) Fr., <italic>Summa Veg.
Scand</italic>.: 383.1849. Figs <xref rid="fig5" ref-type="fig">5</xref>,
<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 citrina</italic> Pers., <italic>Obs</italic>. <italic>Mycol</italic>. 1:
68. 1796: Fr., <italic>Syst</italic>. <italic>Mycol</italic>. 2: 337. 1823.</p></list-item><list-item><p>= <italic>Sphaeria lactea</italic> Fr., K. Svenska VetenskAkad. Handl. II, 37: 141.
1816: Fr., <italic>Syst</italic>. <italic>Mycol</italic>. 2: 337. 1823.</p></list-item><list-item><p>≡ <italic>Hypocrea lactea</italic> (Fr.: Fr.) Fr., <italic>Summa</italic><italic>Veg</italic>. <italic>Scand</italic>.: 383. 1849.</p></list-item></list></p><p><italic>Anamorph</italic>: <italic>Trichoderma lacteum</italic> Bissett [sect.
<italic>Hypocreanum</italic> Bissett], Canad. J. Bot. 69: 2367. 1991.</p><p><italic>Teleomorph</italic>: Stromata effuse, extensive, largest continuous stroma,
140 × 40 mm, smallest continuous stroma, 3 × 2 mm, many stromata
not larger than 25 × 10 mm, solitary to confluent, varying in colour,
sometimes white to light pastel, usually yellow to greyish yellow, sometimes
orange-yellow to brownish yellow (4A4–4B4; 4A6–5A7; 3A4),
KOH<sup>+/–</sup>, reaction variable, usually very weak with slight
browning of stroma; ostiolar canals visible at stroma surface, appearing light
brown, giving rise to the greyish yellow overall appearance of the stroma.
Stroma surface smooth; tissue immediately below the stroma surface and
perithecia of compact to loose pseudoparenchymatous cells consisting of
<italic>textura globulosa</italic> to <italic>t. angularis</italic> with some
compact-vertically arranged hyphae. Perithecia completely immersed except for
a slight protrusion of the ostiolar canals, ≤ 30 μm high (n = 7),
generally widely spaced, compact in some regions, sometimes completely absent
near the margins or regions of extensive stroma growth. Perithecia
ellipsoidal, (263–)275–332(–360) μm long (including the
length of the ostiolar canal, n = 22); width of perithecia near the base
(measured from 3/4 total length of the perithecium)
(99–)122–192(–214) μm (n = 22); length of ostiolar canal
(42–)52–74(–82) μm; width of the ostiolar canal from the
outer perithecial wall to the opposing internal perithecial wall
(29–)34–54(–65) μm (n = 22); wall
KOH<sup>+/–</sup>, reaction variable, weak, usually brownish orange.
Asci cylindrical, (65–)80–102 (–114) ×
(3.7–)6.2–7.2(–9.6) μm (n = 149); tip slightly thickened.
Part-ascospores hyaline, thick-walled, minutely spinulose, dimorphic; distal
part subglobose, sometimes subellipsoidal,
(3.2–)4.1–5.2(–6.3) × (2.9–)
3.7–4.8(–5.7) μm, L/W ratio, (0.7–)0.99-1.2(–1.5)
(n = 166); proximal part, ellipsoidal, sometimes subglobose to ovate,
(3.3–)4.4–6(–7.1) ×
(2.5–)3.3–4.9(–6.3) μm, L/W ratio,
(0.6–)1–1.6(–2.1) (n = 166).</p><p><italic>Anamorph</italic>: Colony covering the surface of a 100 mm diam Petri plate
with PDA in 10 d, not producing concentric rings or radial rays of mycelium; a
layer of cottony aerial mycelium along the entire Petri plate; no distinctive
odour or pigmentation. Conidiophores irregularly branched on long hyphal
elements, usually verticillium-like; phialides in whorls of
3–4(–5), sometimes solitary; phialides subulate,
(8–)14–19 (–26) ×
(2.3–)2.9–3.7(–4.3) μm (n = 61); conidia variable in
size, subglobose to subellipsoidal, (3–)3.3–5.8 (–10.8)
× (3–)3.5–4.3(–5.1) μm (n = 61), some conidia
asymmetric with a truncate base; chlamydospores cylindrical to subglobose when
intercalary and bulbous when terminal, (5–)14-20(–22) ×
(9–)10–15(–18) μm (n = 16). After 10 d, conidia begin to
swell and are more variable in size. Conidiophores not visible at 10 d on SNA
or CMD. Conidiophores developed on CMD and SNA at 15–20 d, primarily
near the point of inoculation. Optimal temperature for growth is 25 °C on
all three media.</p><p><italic>Habitat</italic>: On leaf litter and rich soils, especially soil near
decaying stumps, less commonly on decorticated wood, and moss-covered
bark.</p><p><italic>Known distribution</italic>: Europe, Japan, and North America.</p><p><italic><bold>Neotype</bold></italic> (designated here): <bold>Belgium,</bold> Hestreux near
Eupen: on leaf litter including pine needles, Oct. 1985, W. Gams 4031
(<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>).</p><p><italic>Other specimens examined</italic>: <bold>Austria,</bold> Steiermark,
Lassnitzhöhe: vic. of Graz, elev. 480 m, on leaf litter and pine needles,
1995, H. Voglmayr (BPI 737697; culture G.J.S. 95-96); Niederösterreich,
near Krems at Waldhof, beside Lake Egelsee, on leaf litter of
<italic>Fagus</italic>/<italic>Picea</italic>, 28 Aug. 2000, W. Klofac, comm. W. Jaklitsch
1617 (BPI 748251; culture G.J.S. 00-161). <bold>England,</bold> Devon, Budleigh, on
ground near base of <italic>Picea sitchensis</italic> stump, moss present, 22 Oct.
1989, J. Webster (BPI 1107145; culture G.J.S. 89-145). <bold>Germany,</bold> Pelmer
Wald near Gerolstein, Oct. 1970, W. Gams
(<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>).
<bold>Netherlands,</bold> Baarn (Zandheuvelweg), Aug. 1973, W. Gams
(<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>)<bold>.
Sweden,</bold> in mixed forest between Heina and Saara, on birch bark, 14 Aug.
1979, Lars Fagerström (BPI 744475); Småland, Femsjö, on wood,
date not known, E. M. Fries (UPS, as <italic>Hypocrea</italic> <italic>lactea</italic>);
Femsjö, on dicot leaf, date not known, E. M. Fries (UPS, as <italic>Sphaeria
lactea</italic>), designated here as <bold>lectotype</bold> for <italic>H. lactea</italic>;
Uppland, Arentuna parish, Storvreta, Eastern outskirts, on stump of <italic>Salix
cinerea</italic>, 12 Sep. 1987, N. Lundqvist 17028 (UPS). <bold>USA,</bold> Georgia,
Rabun Co., Chattahoochee Natl. Forest, along Little Creek Rd., near Double
Bridge Creek, 34°59' N, 83°10' W, 17 Oct. 1990, A. Y. Rossman & G.
J. Samuels, G.J.S. 90-137 (BPI 1107189); Kentucky, Laurel Co., Daniel Boone
National Forest, elev. 350 m, Laurel River Lake Recreation Area, Cane Creek
Wildlife Refuge, on decorticated wood, 27 Sep. 1995, G. J. Samuels (BPI
737759; culture G.J.S. 95-183); New York, Oswego County, Vandercamp Trail, on
the moss <italic>Dreplanocladus</italic> <italic>uncinatus</italic>, 2 Oct. 1999, B.E.
Overton, B.E.O. 99-29 (BPI); Warren County, Pack forest North of Warrensburg,
on litter under white pine, 9 Sep. 1963, R.L. Gilbertson 4195 (UPS); North
Carolina, Giles Co., off Bullpen Rd., 35°01' N, 83° 08' W, elev. 3000
ft., 14 Oct. 1990, Y. Doi, A.Y. Rossman & G.J. Samuels, G.J.S. 90-104 (BPI
1107172). Ohio: on <italic>Salix</italic> sp., 3 Oct. 1896, A. P. Morgan No. 46 (BPI
631486).</p><p><italic>Comments</italic>: Gams 4031 (herb. CBS, ex-neotype culture
<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>) is
designated here as the <bold>neotype</bold> specimen of <italic>H. citrina</italic>. This
European specimen is well suited to serve as the neotype because it is yellow,
effuse, and occurs on leaf litter just as Persoon described. <italic>Hypocrea
citrina</italic> is similar in overall appearance to <italic>H. sulphurea</italic>, but
<italic>H. sulphurea</italic> is not as effuse near the stroma margin, is intensely
yellow throughout, and occurs on <italic>Exidia</italic> sp. on twigs, branches, and
trunks.</p><p>A <bold>lectotype</bold> is designated here for <italic>Hypocrea lactea</italic> [Sweden,
Småland, Femsjö (as <italic>Sphaeria lactea</italic>, UPS herb. E.M.
Fries)], assuming that this material was collected before 1822 (otherwise it
would be neotype). <italic>Hypocrea lactea</italic> as lectotypified here is therefore
a synonym of <italic>H. citrina</italic>. Doi
(<xref ref-type="bibr" rid="ref9">1972</xref>) illustrated stroma
characters and part-ascospore measurements in his composite description of
<italic>H. lactea</italic> that are not present in, or indicative of the lectotype
specimen designated here for <italic>H. lactea s.s.</italic> Doi's (1979) annotation
of Fries's historical specimen of <italic>H. lactea</italic> (cited in his specimens
examined) listed part-ascospore measurements as follows: distal part-spores
3.5–4.5 × 3.4–3.9 μm, proximal part-spores 4.5–5.5
× 3.2–3.5 μm. These measurements differ substantially from the
measurements given by Doi
(<xref ref-type="bibr" rid="ref9">1972</xref>) in his composite
description of <italic>H. lactea</italic>; distal part-ascospores 4.5–5.2
× 4.5–4.8 μm, proximal part-spores, 5.5–7.1 ×
4.2–4.4 μm. The Japanese specimens cited by Doi under the name <italic>H.
lactea</italic> likely represent an undescribed species from Japan, especially
given that Doi illustrated hyphal protrusions near the stroma surface, which
are not indicative of <italic>H. lactea</italic>.</p></list-item><list-item><p><bold><italic>Hypocrea pulvinata</italic></bold> Fuckel, Jahrb. Nassau. Ver. Naturk.
23–24: 185. 1870 [“1869”]. Figs
<xref rid="fig9" ref-type="fig">9</xref>,
<xref rid="fig10" ref-type="fig">10</xref>,
<xref rid="fig11" ref-type="fig">11</xref>.</p><p><list list-type="simple"><list-item><p>= <italic>Hypocrea citrina</italic> * <italic>fungicola</italic> P. Karst., <italic>Mycol.
Fenn</italic>. 2: 204. 1873.</p></list-item><list-item><p>≡ <italic>Hypocrea karsteniana</italic> Niessl in Rehm. Ascomyceten, Hedwigia
22: 53 [Apr.] 1883.</p></list-item><list-item><p>≡ <italic>Hypocrea fungicola</italic> (P. Karst.) Sacc., Syll. Fung. 2: 528
[13 June] 1883.</p></list-item><list-item><p>≡ <italic>Protocrea fungicola</italic> (P. Karst) Lar.N. Vassiljeva,
<italic>Nizshie Rasteniya, Griby i Mokhoobraznye Dal'nego Vostoka Rossii,
Griby.</italic> Tom 4. <italic>Pirenomitsety i Lokuloaskomitsety</italic>(Sankt-Peterburg): 162. 1998.</p></list-item><list-item><p>= <italic>Hypocrea citrina</italic> (Fr.) Fr. var. <italic>citrina sensu</italic> Cunham,
Mycologia 61: 318. 1969.</p></list-item></list></p><p><italic>Anamorph</italic>: <italic>Trichoderma</italic> sp. [sect.
<italic>Hypocreanum</italic>].</p><p><italic>Teleomorph</italic>: Stromata effuse to subpulvinate, extensive, solitary
to confluent, largest continuous stroma 60 × 20 mm, smallest continuous
stroma 3 × 2 mm, varying in colour, orangish yellow to greyish orange,
sometimes brownish yellow to golden-yellow, or brown-yellow (4B7–4B6;
5C7–5B7; 5C5); KOH<sup>+</sup>, reaction variable, sometimes weak,
generally with stromata becoming orange or red; ostiolar openings visible from
stroma surface. Warted hairs at surface of stromata, tissue immediately below
the stroma surface formed of compact to loose pseudoparenchymatous cells,
<italic>textura globulosa</italic> to <italic>t. angularis</italic>. Perithecia numerous and
completely immersed in the stromata, generally widely spaced but compact near
the centre, ellipsoidal, (189–) 206–246(–275) μm long
(including length of ostiolar canal, n = 28); width of perithecia near the
base (3/4 total length of perithecium) (96–)105–161(–186)
μm (n = 28); length of ostiolar canal (36–)48–72(–82)
μm; width of ostiolar canal (29–)43–63(–72) μm (n =
28); perithecial wall KOH<sup>+</sup>, reaction variable, usually brownish
orange. Asci cylindrical, (44–)66–90(–115) ×
(2.8–)3.6–5.2(–7.9) μm (n = 321), tip slightly thickened.
Part-ascospores hyaline, thin-walled, smooth to minutely spinulose, generally
monomorphic and longer than broad; distal part ellipsoidal, sometimes
subglobose, (2.8–)3.7–4.7(–6.0) ×
(2.3–)2.9–3.7(–4.6) μm, L/W ratio
(0.8–)1.1–1.5(–2.2) (n = 335); proximal part, ellipsoidal,
sometimes subglobose, (2.8–)3.8–5 (–6.7) ×
(2.3–)2.8-3.8(–4.7) μm, L/W ratio,
(0.8–)1.1–1.6(–2.2) (n = 335).</p><p><italic>Anamorph</italic>: Colonies covering a 100 mm diam Petri plate with PDA in
10 d; producing concentric growth pattern with radial rays; small white
cottony aggregates near the edge of the Petri dish composed of mycelium and
acremonium-like conidiophores, no aerial conidiophores; stromata and
perithecia produced near the edge of the plate; reddish brown pigment (9E8)
produced near the centre of the plate and brown pigment (7E8) near the edge of
the plate; no odour. Colonies covering a 100 mm diam Petri plate with SNA also
in 10 d; forming a thin layer with small white cottony aggregates of
conidiophores (at random places on the plate); no aerial mycelium and only a
few scattered conidiophores with conidia; no pigment production, no odour.
Conidiophores developed and started producing conidia on SNA at 10 d. Colonies
covering a 100 mm diam Petri plate with CMD in 10 d; forming a thin layer with
small white cottony aggregates near the edge of the Petri plate; trace of
brown pigment (7E8) near the inoculum; no odour. Conidiophores developing on
CMD, only a few producing conidia. Optimal temperature for growth on all 3
media is 25 °C.</p><p>Measurements of micro-morphological characters were the same on all three
media. A description based on measurements from all three media at 10 d
follows: conidiophores variable in branching pattern and phialide number;
phialides generally solitary, subulate, variable in length,
(7–)24–44(–50) × (2.7–)3.6–4.6
(–5.3) μm (n = 32); below the phialides chlamydospore-like hyphal
thickenings sometimes observed; conidia variable in size, many asymmetric,
having a truncate base, subglobose to subellipsoidal, some bulb-shaped,
(2.7–)4.1–7.1(–11) ×
(2.1–)3.3–5.6(–9) μm (n = 92). After 10 d conidia began
to swell and were more variable in size.</p><p><italic>Habitat</italic>: Found on a variety of polypores, including <italic>Laetiporus
sulphureus, Fomitopsis pinicola, Piptoporus betulinus</italic>, and
<italic>Ganoderma</italic> spp.</p><p><italic>Known distribution</italic>: Europe, Japan, and North America.</p><p><italic><bold>Lectotype</bold></italic> (designated here): <bold>Germany,</bold> Geis,
Hattenheimer Wald, on <italic>Polyporus sulphureus</italic> (= <italic>Laetiporus</italic><italic>sulphureus</italic>), L. Fuckel, autumn, No. 876 (FH).</p><p><italic>Other specimens examined</italic>: <bold>Austria,</bold> Wachau District, Krems
an der Donau, Mosingraben, North of Spitz in mixed forest, elev. 300 m, on
undetermined polypore, 2 June 1996, I. Krisai-Greilhuber (BPI 744499; culture
G.J.S. 96-142); Waldviertel, Virgin Forest of Dobra, on <italic>Fomitopsis
pinicola</italic>, 10 Oct. 1995, H. Voglmayr (BPI 737803; culture G.J.S. 95-220).
<bold>Finland,</bold> Locality unknown, Fungi Fennici Exs. 264, identified as <italic>H.
citrina</italic> f. <italic>fungicola</italic> (K); Wasa (= Vassa), on polypore, May 1869,
P.A. Karsten s.n. (H, herb. Karsten 4479, as <italic>H. citrina</italic> f.
<italic>fungicola</italic>); Tammela, Mustiala, on polypore, 25 Sep. 1890, P. A.
Karsten s.n. (H, herb. Karsten 4475); Satakunta Tyrvis (= Vammala,
Tyrvää), on polypore, date unknown, P.A. Karsten s. n. (H, herb.
Karsten 4478; syntype of <italic>H. citrina</italic> f. <italic>fungicola</italic>); Karelia
Keretina, on <italic>Polyporus</italic> (<italic>Piptoporus</italic>) <italic>betulinus</italic>, 9
Aug. 1861, P.A. Karsten s.n. (H, herb. Karsten 4477, as <italic>H. citrina</italic> f.
<italic>fungicola</italic>; syntype). <bold>Germany,</bold> Geis, on “<italic>Polyporus
resinosus”</italic>, substrate determined as <italic>Laetiporus sulphureus</italic>by E. Parmasto (Oct. 1999), Fuckel, Fungi Rhenani 2467 (K 61843); Saarland,
Naturpark Saar-Hundsrück, vic. Ritzenberg, 90°40' N, 7°0' W,
elev. 600–650 m, on <italic>Ganoderma</italic> sp., 12 Oct. 1998, G. J. Samuels
& H. J. Schroers, G.J.S. 98-104 (BPI 746114); BPI 746115; culture G.J.S.
98-108. <bold>Great Britain,</bold> Yorks., Wrathe Woods, Pately Bridge, on
<italic>Piptoporus betulinus</italic>, 25 Sep. 1955, C. Booth, IMI 61207 (BPI 631478).
<bold>Luxembourg,</bold> Nassebesh, Hachy, on <italic>Piptoporus betulinus</italic>, 19 Sep.
1996, Vanhalle S. Mull 40055 (BPI 744722; culture G.J.S. 96-283).
<bold>Poland,</bold> Silesia, Karlsbrunn, on <italic>Polyporus pinicola</italic> (=
<italic>Fomitopsis pinicola</italic>, conf. E. Parmasto), Aug. 1882, von Niessl, Rehm
Ascomyceten 678 (K, 61844). <bold>Sweden,</bold> on polypore, E.M. Fries,
Scleromyceti Sueciae 31 (K). <bold>U.S.A.,</bold> New Hampshire, Chocorua, on
<italic>Fomitopsis</italic> <italic>pinicola</italic>, 16 July 1906, W.G. Farlow No. 34,
identified as <italic>H. fungicola</italic> (K); New York, Franklin County, Little
Green Pond, vic. Paul Smith College, on <italic>Ganoderma</italic> sp., 17 Aug. 1991,
R. Lowen R.L. 886 (BPI 112828; culture G.J.S. 91-55A); Oneida County, on
<italic>Fomitopsis pinicola</italic>, Barrie Overton, B.E.O. 99-34 (BPI); Utah, Beaver
County, Tushar Mountains, Fishlake National Forest, on <italic>Fomitopsis
pinicola</italic>, 31 Aug. 1994, C.T. Rogerson (BPI 737831; culture G.J.S. 94-20);
Washington, Olympia National Park, Twin Creek, on <italic>Fomitopsis
pinicola</italic>, 18 Oct. 1992, H. Burdsall, H.B. 14889 (BPI 802876; culture
G.J.S. 92-127); Olympia National Park, Twin Creek, on <italic>Fomitopsis
pinicola</italic>, 18 Oct. 1992, H. Burdsall, H.B. 14910 (BPI 802877; culture
G.J.S. 92-128).</p></list-item><list-item><p><bold><italic>Hypocrea americana</italic></bold> (Canham) Overton, <bold>stat. nov</bold>.
MycoBank <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB501053&link_type=MB">MB501053</ext-link>.
Figs <xref rid="fig12" ref-type="fig">12</xref>,
<xref rid="fig13" ref-type="fig">13</xref>,
<xref rid="fig14" ref-type="fig">14</xref>.</p><p><list list-type="simple"><list-item><p>≡ <italic>Hypocrea citrina</italic> (Fr.) Fr. var. <italic>americana</italic> Canham,
Mycologia 61: 320. 1969.</p></list-item></list></p><p><italic>Anamorph</italic>: <italic>Trichoderma</italic> sp. [sect.
<italic>Hypocreanum</italic>].</p><p><italic>Teleomorph</italic>: Stromata effuse to subpulvinate, solitary to
confluent, extensive, largest continuous stroma 50 × 20 mm, smallest
continuous stroma 3 × 2 mm, varying in colour, usually orange to
brownish yellow, or orangish yellow to greyish yellow (5B8–5C8;
4B7–4B6); KOH<sup>+</sup>, reaction variable, generally with stromata
becoming dark orange or red; perithecial necks or openings barely visible from
stroma surface. Warted hairs at stroma surface; tissue immediately below the
stroma surface formed of compact to loose pseudoparenchymatous cells composed
of <italic>textura globulosa</italic> to <italic>t. angularis</italic>. Perithecia numerous
and completely immersed in the stromata, generally widely spaced but compact
near the centre of the stroma, ellipsoidal, (149–)167–199
(–213) μm long (including the length of the ostiolar canal, n = 15);
width of perithecium near the base, measured from 3/4 the total length of the
perithecium, (73–)80–106 (–119) μm (n = 15); length of
ostiolar canal (44–)54–74 (–84) μm; width of ostiolar
canal (41–)45–56(–62); wall KOH<sup>+</sup>, reaction
variable, usually brownish orange or red. Asci cylindrical,
(49–)59–79(–106) × (3.3–)4–5(–6.2)
μm (n = 167), tip slightly thickened. Part-ascospores hyaline, thin-walled,
smooth, monomorphic, usually subglobose to rhomboidal by mutual pressure,
broader than long; distal part subglobose to rhomboidal, (1.7–)
2.7–3.7(–6.2) × (2.4–)3.2–4.2(–5.3) μm,
L/W ratio (0.5–) 0.84–0.89(–1.5) (n = 212); proximal part
subglobose to rhomboidal, (1.8–)2.6–3.8(–5.7) ×
(2.5–)3.2–4.2(–5.4), L/W ratio
(0.5–)0.82–0.86(–1.4) (n = 212).</p><p><italic>Anamorph</italic>: Colonies covering a 100 mm diam Petri plate with PDA in
10 d; mycelium growing tightly pressed to the surface, appearing wet, no
aerial mycelium; conidiophores produced in cottony tufts concentrated near the
edge of the plate, tufts thicker and rising further from the agar surface than
on CMD; stromata and perithecia emerging from several regions where
conidiophores are found in the greatest number; brownish orange (7C8) to
reddish brown (8E8) pigment diffusing in the medium, pigments visible within
the hyphae. Colonies covering a 100 mm diam Petri plate with SNA in 10 d;
mycelium thin, watery in appearance; conidiophores scattered, conidial
production poor or absent. Colonies covering a 100 mm diam Petri plate with
CMD in 10 d; mycelium tightly pressed to the surface, appearance watery, no
aerial mycelium; conidiophores produced in cottony aggregates and tufts
concentrated near the edge of the plate; brownish orange (7C8) to light brown
pigment (7D8) diffusing into the medium. Optimal temperature for growth on all
3 media is 25°C.</p><p>Measurements of micro-morphological characters were the same on all three
media. A description based on measurements from all three media at 10 d
follows: conidiophores variable in branching pattern and phialide number;
phialides generally solitary, subulate, variable in length,
(12–)20–40(–56) ×
(1.8–)3.3–5.1(–6.4) μm (n = 41), with chlamydospore-like
thickenings just below the phialides on CMD and PDA; conidia variable in size
and shape, typically subglobose to subellipsoidal, sometimes bulb-shaped, many
asymmetric, having a truncate base, (3.2–)3.6–6.8(–10.2)
× (2.6–)3.2–5.9 (–8.8) μm (n = 43). After 10 d
conidia begin to swell, becoming larger and more variable in size.</p><p><italic>Habitat</italic>: On <italic>Fomitopsis pinicola</italic> and <italic>Piptoporus
betulinus</italic>.</p><p><italic>Known distribution</italic>: North America.</p><p><italic>Holotype</italic>: <bold>USA,</bold> New York, Madison Co., Nelson Swamp, on
<italic>F</italic>. <italic>pinicola</italic>, 9 Oct. 1949, C.T. Rogerson 3284 &
Huttleston (NY ex CUP 38045, ex-type culture ATCC 18574).</p><p><italic>Other specimens examined</italic>: <bold>U.S.A.,</bold> Arizona, Apache Co.,
White Mountains, 19.1 miles from forest boundary, on <italic>F</italic>.
<italic>pinicola</italic>, 21 Aug. 1994, C.T. Rogerson (BPI 737849; culture G.J.S.
94-79); Graham Co., Pin leno Mt., Coronado National Forest, Rt. 366, 4.1 miles
south of Riggs Lake, on <italic>F. pinicola</italic>, 15 Aug. 1996, C.T. Rogerson (BPI
744432; G.J.S. 96-191); New Hampshire, East Monroe, on <italic>Piptoporus
betulinus</italic>, 22 Aug. 1937, C.L. Shear (BPI 631477); New Mexico, Upper
Fernando de Taos, forest service map, T25n R15e S15, on <italic>Fomes</italic> sp., 15
Aug. 1992, R. Lowen 1003 (BPI 802838; culture G.J.S. 92-93); New York, Oswego
Co., Vandercamp Trail, on <italic>F</italic>. <italic>pinicola</italic>, 2 Oct. 1999, B.E.
Overton, B.E.O. 99-30 (BPI); also B.E.O. 99-32 and B.E.O. 99-33 (all BPI);
Ulster County, Clinton dale Swamp, on <italic>P</italic>. <italic>betulinus</italic>, 11 Oct.
1963, S. J. Smith, R. Detroit, & C. T. Rogerson (NY; <bold>paratype</bold>).</p><p>Comments: <italic>Hypocrea americana</italic> is easily distinguished from <italic>H.
pulvinata</italic> by size and shape of the ascospores and degree of visibility of
the ostiolar openings. The ascospores of <italic>H. americana</italic> are globose to
rhomboidal with width greater than the length. In <italic>H. pulvinata</italic>, the
ascospores are ellipsoidal, the length exceeding the width. The ostiolar
openings in <italic>H. pulvinata</italic> are visible and the stroma surface has less
of an overall rugulose appearance than in <italic>H. americana</italic>. In the
specimen BPI 802838, both <italic>H. pulvinata</italic> and <italic>H. americana</italic>occur on the same host.</p></list-item><list-item><p><bold><italic>Hypocrea protopulvinata</italic></bold> Yoshim. Doi, Bull. Natl. Sci. Mus.
15: 695. 1972. Figs <xref rid="fig15" ref-type="fig">15</xref>,
<xref rid="fig16" ref-type="fig">16</xref>.</p><p><italic>Anamorph</italic>: <italic>Trichoderma</italic> sp. [sect.
<italic>Hypocreanum</italic>].</p><p><italic>Teleomorph</italic>: Stromata effuse, sometimes subpulvinate, solitary to
confluent, extensive, varying in colour, usually orangish yellow to light
brown, young stromata white, becoming pale yellow where perithecia are
developing (4B6–6D7; 4A3); KOH<sup>+/–</sup>, reaction variable,
weak, generally with stromata becoming darker orange; largest continuous
stroma 22 × 15 mm, smallest continuous stroma 5 × 3 mm; ostiolar
openings visible from stroma surface. Hyphal protrusions at stroma surface
subglobose to pyriform, slightly roughened, not appearing warted like in
<italic>H. pulvinata</italic>. Tissue immediately below the stroma surface variable;
cells adjacent to ostiolar canals composed of rather thick-walled <italic>t.
angularis</italic>; cells between perithecia of <italic>t. globulosa</italic> with some
<italic>t. intricata</italic> intermixed; cells near the base of the perithecia
composed of tightly packed irregular pseudoparenchymatous tissue in between
<italic>t. globulosa, t. intricata</italic>, and <italic>t. angularis</italic>. Perithecia
numerous near the stroma centre, scarce near the margin, generally widely
spaced, subglobose to ellipsoidal, (151–)180–286(–310) μm
long (including the length of the ostiolar canal, n = 11); width of perithecia
near the base (99–)106–153(–176) μm, n = 11; length of
ostiolar canal (33–)33–57(–68) μm; width of ostiolar
canal (27–)30–46(–53) μm, n = 11; wall
KOH<sup>+/–</sup>, reaction variable, weak, usually brownish orange.
Asci cylindrical, (45–)56–72(–76) ×
(3.7–)4–5(–5.4) μm (n = 52), tip slightly thickened.
Part-ascospores hyaline, thin-walled, smooth, sometimes minutely spinulose,
dimorphic; distal part subglobose, (3–)3.3–3.9(–4.3) ×
(2.6–)3–3.5(–4.2) μm, L/W ratio,
(0.8–)0.99–1.2 (–1.5) (n = 52); proximal part ellipsoidal,
sometimes subglobose, (3.2–)3.6–4.2(–4.8) ×
(2.4–)2.7–3.3(–3.8), L/W ratio
(0.9–)1.1–1.5(–1.8) (n = 52).</p><p><italic>Anamorph</italic>: Colonies growing across the surface of a 100 mm diam
Petri plate with PDA in 10 d; radial growth with radial rays, mycelium tightly
pressed to the surface, sometimes with web-like sectors of rapidly growing
mycelium, no aerial mycelium; conidiophores produced near the edge of the
colonies in tufts; several large aggregates appear to be stroma initials;
other large aggregates consist of clustered conidiophores; agar with a
yellowish tint from the reverse, brown pigment (5D7) being produced near the
edge of the plate and the agar plugs; sometimes odour (fruity of sour apples)
produced when plates are tightly sealed; stromata produced 25–30 d
following inoculation in sealed plates of the ex-type strain. Colonies growing
across the surface of a 100 mm diam Petri plate with SNA in 10 d; mycelium
web-like, random tufts of conidiophores arising from the agar surface, very
few mature conidiophores; no pigment production. Colonies growing across the
surface of a 100 mm diam Petri plate with CMD in 10 d; mycelium tightly
pressed to the surface, appearing wet, no aerial mycelium; conidiophores
produced near the edge of the plate, very rarely in tufts arising from the
surface, very few mature conidiophores. Optimal temperature for growth on all
3 media is 25–30 °C with cultures growing ≤ 3 mm at 35
°C.</p><p>Measurements of micro-morphological characters were the same on all three
media. A description based on measurements from all three media at 10 d
follows: conidiophores irregular, acremonium-like, variable in branching
pattern and phialide number; phialides generally solitary, subulate, variable
in length, (8–)20–38(–49) ×
(3–)3.7–4.7(–6) μm (n = 54); chlamydospore-like swellings
below the phialides not as prominent on PDA and CMD; conidia variable in size
and shape, generally subellipsoidal, some subglobose to subellipsoidal, most
conidia have a well-developed truncate base,
(4–)5.9–9.9(–16) × (2.8–)4–7(–9.5)
μm (n = 76); chlamydospores not observed after 10 d. After 10 d conidia
begin to swell and are more variable in size.</p><p><italic>Habitat</italic>: On <italic>Fomitopsis pinicola</italic> and unidentified
polypores.</p><p><italic>Known distribution</italic>: Japan and North America.</p><p><italic>Isotype</italic><bold><italic>:</italic></bold> <bold>Japan,</bold> Chiba Pref., Fudagou Mt.,
on unidentified polypore, 24 Oct. 1967, Y. Doi (NY ex TNS.D-365 =
TNS-F-223421; ex-type culture
<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>).</p><p><italic>Other specimens examined</italic>: <bold>U.S.A.,</bold> Maryland, on unidentified
polypore, autumn 2000, Kadri Põldmaa, K.P. 00-56 (BPI); New York,
Cranberry Lake, Suckers Trail, on unidentified polypore, 15 Sep. 2001, B.E.
Overton & D. Geiser, B.E.O. 00-01 (BPI); Ohio, on unidentified polypore,
11 Nov. 1883, A.P. Morgan No. 29 (BPI 631312; specimen degraded).</p><p><italic>Comments</italic>: <italic>Hypocrea protopulvinata</italic> is easily distinguished
from <italic>H. pulvinata</italic> and <italic>H. americana</italic> because of its dimorphic
ascospores and more effuse stromata with roughened, not warted, hyphal
protuberances. The subglobose distal part-spores of <italic>H. protopulvinata</italic>are similar in size to the monomorphic subglobose part-spores found in <italic>H.
americana</italic>: (3–)3.3–3.9(–4.3) ×
(2.6–)3–3.5(–4.2) μm in <italic>H. protopulvinata</italic> versus
(1.7–)2.7–3.7(–6.2) ×
(2.4–)3.2–4.2(–5.3) μm in <italic>H. americana</italic>; In
addition, the ellipsoidal proximal part-spores of <italic>H. protopulvinata</italic>are similar in size to the monomorphic ellipsoidal part-spores found in <italic>H.
pulvinata</italic>: (3.2–)3.6–4.2(–4.8) ×
(2.4–)2.7–3.3(–3.8) in <italic>H. protopulvinata</italic> versus
(2.8–)3.8–5(–6.7) ×
(2.3–)2.8–3.8(–4.7) in <italic>H. pulvinata</italic>. All three of
these species have thin-walled part-ascospores that are smooth to minutely
spinulose, characters not common in <italic>Hypocrea</italic>.</p></list-item><list-item><p><bold><italic>Hypocrea platypulvinata</italic></bold> Yoshim. Doi, Bull. Natl. Sci. Mus.
15: 698. 1972.</p><p><italic>Anamorph</italic>: <italic>Trichoderma</italic> sp. [sect.
<italic>Hypocreanum</italic>].</p><p><italic>Teleomorph</italic>: Stromata in aggregates or scattered, disciform,
attached to decayed logs via a narrow base, deep yellow-ochre to ochre-brown,
2–6 mm diam, 0.8–1.6 mm thick, cells at surface of stromata
roughened; perithecia ellipsoid, 200–240 μm in length;
part-ascospores hyaline, warted, generally monomorphic, globose, 2.1–2.3
μm diam or sometimes obovate ellipsoid, 2.0–3.0 μm long
(<xref ref-type="bibr" rid="ref9">Doi 1972</xref>).</p><p><italic>Anamorph</italic>: Conidiophores acremonium-like; phialides subulate toward
the apices, 15–35 × 1.5–2.0 μm; conidia hyaline, smooth,
cylindrical to obovate-ellipsoid, with a truncate base, 4.0–14 ×
2.3–5.1 μm; chlamydospore-like, thick-walled cells often produced,
2–12 × 6–8 μm in culture
(<xref ref-type="bibr" rid="ref9">Doi 1972</xref>).</p><p><italic>Habitat</italic>: On decayed logs of broad-leaved trees.</p><p><italic>Known distribution</italic>: Japan.</p><p><italic>Specimens examined</italic>: Type material not available for
examination.</p></list-item><list-item><p><bold><italic>Hypocrea megalocitrina</italic></bold> Yoshim. Doi, Bull. Natl. Sci. Mus.
15: 669. 1972. Figs <xref rid="fig17" ref-type="fig">17</xref>,
<xref rid="fig18" ref-type="fig">18</xref>.</p><p><italic>Anamorph</italic>: <italic>Trichoderma</italic> sp. [sect.
<italic>Hypocreanum</italic>].</p><p><italic>Teleomorph</italic>: Stromata effuse, solitary to confluent, extensive,
largest continuous stroma, 40 × 30 mm, smallest continuous stroma 3
× 2 mm, varying in colour, usually light brown (6D7);
KOH<sup>–</sup>; surface of stromata appearing roughened due to ostiolar
canals projecting from the stroma surface, which are covered by a thin layer
of <italic>t. intricata</italic>. Tissue immediately below the stroma surface formed
of compact to loose <italic>t. intricata</italic> and sporadic pseudoparenchymatous
cells varying from <italic>textura globulosa</italic> to <italic>t. angularis</italic>.
Perithecia numerous, generally widely spaced but compact near the centre of
the stroma, ellipsoidal-elongate, (207–)240–320 (–345) μm
high (including the length of the ostiolar canal, n = 24); width of perithecia
near the base, measured from 3/4 the total length of each perithecium,
(113–) 127–181(–208) μm (n = 24); length of ostiolar
canal (54–)62–94(–130) μm (n = 24); width of ostiolar
canal (54–)57–75(–84) (n = 24); wall KOH<sup>–</sup>.
Asci cylindrical, (66–)79–97(–118) ×
(4.7–)5.3–6.6(–7.4) μm (n = 93), tip slightly thickened.
Part-ascospores hyaline, thick-walled, nodulose, dimorphic; distal part
subellipsoidal to conical, (3.2–)4.5–5.8(–6.6) ×
(3.0–)3.8–5(–6) μm, L/W ratio
(0.9–)1–1.3(–1.9) (n = 121); proximal part ellipsoidal,
rarely subellipsoidal, sometimes appearing thimble-shaped or conical,
(3.7–)4.6–5.9(–7) × (3.1–)
3.7–4.8(–5.8), L/W ratio (0.8–)1.0–1.4(–1.8) (n
= 121).</p><p><italic>Anamorph</italic>: Colonies on n PDA covering the surface of a 100 mm diam
Petri plate in 10 d, producing an extensive layer of aerial mycelium;
conidiophores concentrated near the agar plug, some areas with thicker layers
of aerial mycelium than others; conidiophores in dense clusters near the agar
plug, but also on thin layers of aerial mycelium near the edge of the plate;
no diffusible pigment observed in the agar; conidiophores typically
verticillately branched; phialides slender, subulate, typically in whorls of
3–5, but also solitary; phialides with conidia appearing light yellow
when compacted in the aerial mycelium, (11–)15–24(–30)
× (2–)2.3–3.4(–4) μm; conidia hyaline, smooth,
subglobose to ellipsoidal, variable in size and shape, sometimes with a
truncate base, (3.8–)4–6.5(–9) ×
(3.3–)3.4–4(–4.3) μm; chlamydospores not observed at 10
d. Cultures on SNA or CMD did not produce conidiophores within 10 d.</p><p><italic>Habitat</italic>: On decayed trunks of broadleaf trees including
<italic>Acer</italic> sp.</p><p><italic>Known distribution</italic>: Japan and North America.</p><p><italic>Isotype</italic>: <bold>Japan,</bold> Mie, Hirkura Forestry Experimental Station,
7 Sep. 1965, Doi (NY ex TNS-F-223220).</p><p><italic>Other specimens examined</italic>: <bold>U.S.A.,</bold> Maryland, Howard Co. and
Caroll Co. border, Patapsco Valley State Park, N. of Ellicot city, on wood, 14
Oct. 1999, probably <italic>Acer</italic> sp., B.E. Overton, B.E.O. 99-42 (BPI); North
Carolina, Macon Co., Glen Falls lower trail, Nantahala National Forest, 12
July 2000, L. Grand & C. Vernia (BPI, culture B.E.O. 00-09).</p></list-item><list-item><p><bold><italic>Hypocrea aurantiistroma</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=MB501054&link_type=MB">MB501054</ext-link>. Figs
<xref rid="fig19" ref-type="fig">19</xref>,
<xref rid="fig20" ref-type="fig">20</xref>.</p><p><italic>Anamorph</italic>: Unknown.</p><p>Stromata effusa, solitaria vel confluentia, ad 8 × 5 mm,
griseo-aurantia, ostiolis protrudentibus verrucosa. Ascosporae hyalinae,
crassitunicatae, exigue nodulosae, partes quasi monomorphicae; pars distalis
quasi conica, (3.6–)4.5–5(–5.9) ×
(2.5–)3.3–4(–4.6) μm, pars proxima plus minusve
ellipsoidea, (3.6–)4.2–5.4(–6.4) ×
(3.1–)3.3-4(–4.6) μm. Anamorphosis ignota.</p><p><italic>Teleomorph</italic>: Stromata effuse, solitary to confluent, extensive,
largest continuous stroma, 8 × 5 mm, smallest continuous stroma, 2
× 2 mm, greyish orange (6B4); KOH<sup>–</sup>; surface of stromata
appearing roughened because of ostiolar canals projecting from stroma surface;
projecting ostiolar canals covered by a thin layer of <italic>t. intricata</italic>.
Tissue immediately below the stroma surface formed of compact to loose <italic>t.
intricata</italic> and sporadic pseudoparenchymatous cells, typically <italic>textura
globulosa</italic>. Perithecia numerous and completely immersed in stromata,
generally widely spaced but compact near the centre of the stroma,
ellipsoidal-elongate, (207–)240–258(–283) μm high
(including the length of the ostiolar canal, n = 10); width of the perithecium
near the base, measured from 3/4 the total length of each perithecium,
(113–)135–149(–193) μm, n = 10; length of ostiolar canal
(54–)64–90(–99) μm, n = 10; width of ostiolar canal
(67–)69–79(–84) (n = 10); wall KOH<sup>–</sup>. Asci
cylindrical, (66–)80–98(–104) × (4.7–)
5–5.8(–6.4) μm (n = 30), tip slightly thickened;
part-ascospores uniseriate. Part-ascospores, hyaline, thick-walled, minutely
nodulose, slightly dimorphic, almost appearing monomorphic; distal part
subellipsoidal, slightly conical, (3.6–)4.5–5(–5.9) ×
(2.5–)3.3–4(–4.6) μm, L/W ratio,
(0.9–)1–1.4(–1.9) (n = 30); proximal part ellipsoidal,
rarely subellipsoidal, slightly conical, (3.6–)4.2–5.4(–6.4)
× (3.1–)3.3-4(–4.6) μm, L/W ratio,
(0.8–)1.1–1.5(–1.7) (n = 30).</p><p><italic>Habitat</italic>: On bark of <italic>Pinus strobus</italic>.</p><p><italic>Known distribution</italic>: North America, Japan (?).</p><p><italic>Holotype</italic>: <bold>U.S.A.,</bold> North Carolina, Macon, Clear Creek road,
35°00' N, 83°15' W, on <italic>Pinus strobus</italic>, 16 Oct. 1990, Y. Doi,
A. Y. Rossman, G.J. Samuels, (BPI 1107193<bold>;</bold> Doi-68 in TNS, isotype).</p><p><italic>Comments</italic>: <italic>Hypocrea aurantiistroma</italic> can be distinguished
from <italic>H. megalocitrina</italic> by its greyish orange stroma and minutely
nodulose ascospores. This species is similar in overall appearance to <italic>H.
fulva sensu</italic> Yoshim. Doi, which differs substantially from <italic>H. fulva
s.str.</italic> Japanese specimens included under the name <italic>H. fulva</italic> were
not available for study. It is likely that specimens from Japan included under
the name <italic>H. fulva</italic> by Doi represent an undescribed sister species
close to <italic>H. aurantiistroma</italic> or possibly, the same species.</p></list-item><list-item><p><bold><italic>Hypocrea fulva</italic></bold> Penz. & Sacc., <italic>Malpighia</italic> 11:
520. 1897. <xref rid="fig21" ref-type="fig">Fig. 21</xref>.</p><p><italic>Anamorph</italic>: unknown.</p><p><italic>Teleomorph</italic>: Stromata subpulvinate to pulvinate, discrete but
sometimes confluent, largest stroma composed of confluent fructifications, 1.5
× 1.2 mm, smallest continuous stroma 0.5 × 0.5 mm, stromata
generally discrete, 0.55–0.9 mm diam (n = 26), brownish yellow in
lighter regions to yellowish brown in darker regions (5C8–5E8); tissue
immediately below the stroma surface formed of a thin layer (3–4 cells
thick) of loose to compact pseudoparenchymatous cells, typically <italic>textura
globulosa</italic> to <italic>t. angularis</italic>, stromata otherwise composed of
vertically arranged <italic>t. intricata</italic>. Perithecia completely immersed in
the stromata, generally widely spaced, ellipsoidal,
(283–)290–340(–345) μm high (including the length of the
ostiolar canal (n = 4); width of perithecia near the base, measured from 3/4
the total length of each perithecium, (200–)210–250(–258)
μm (n = 3); length of ostiolar canal (63–)64–84(–88)
μm (n = 4). Asci cylindrical, (110–)125–140(–162) ×
(5.6–)6.3–8.0(–9.6) μm (n = 26), tip slightly thickened.
Part-ascospores hyaline, thick-walled, nodulose, slightly dimorphic; distal
part subellipsoidal, slightly conical, (3.9–)4.5–5.5(–6.2)
× (2.8–)3.2–3.8(–4.3) μm, L/W ratio,
(1.1–)1.2–1.6(–2.0) (n = 30); proximal part ellipsoidal,
rarely conical, (4.3–)4.6–5.6(–6.5) × (2.8–)
3.0-3.6(–4.1), L/W ratio, (1.2–)1.3–1.7(–1.9) (n =
30).</p><p><italic>Syntypes examined</italic>: Indonesia, Tjibodas, on herbaceous stem, 104
and 410 (PAD).</p><p><italic>Comments</italic>: <italic>Hypocrea fulva</italic> is very similar to <italic>H.
aurantiistroma</italic> and <italic>H. megalocitrina</italic> as it has ostiolar canals
projecting from the stroma surface, and nodulose part-ascospores. <italic>Hypocrea
fulva</italic> is distinguished by its yellow-brown pulvinate stroma, which has a
thin layer of pseudoparenchymatous tissue, and its slightly conical
part-ascospores. The part-ascospores of <italic>H. megalocitrina</italic> are much
more conical and the nodules more developed, while the part-ascospores of
<italic>H. aurantiistroma</italic> are even less conical with nodules that are present
but not as distinct. The syntypes of <italic>H. fulva</italic> differ morphologically
from the illustrations provided by Doi
(<xref ref-type="bibr" rid="ref9">1972</xref>) under the name <italic>H.
fulva</italic> both in colour and the presence of a well-defined layer of
pseudoparenchymatous tissue at the stroma surface. <italic>Hypocrea fulva
sensu</italic> Doi is likely an undescribed species closely related to <italic>H.
aurantiistroma</italic>. Nodulose ascospores are unusual in Subsection
<italic>Citrinae</italic> as all other species Doi included in <italic>Citrinae</italic> have
smooth to minutely spinulose part-ascospores.</p></list-item><list-item><p><bold><italic>Hypocrea mikurajimensis</italic></bold> Yoshim. Doi, Mem. Natl. Sci. Mus.,
Tokyo 37: 113. 2001.</p><p><italic>Anamorph</italic>: <italic>Trichoderma</italic> sp. [sect.
<italic>Hypocreanum</italic>].</p><p><italic>Teleomorph</italic>: Stromata pulvinate, gregarious or dispersed, sometimes
fused together to form larger and irregular stroma masses, wax-yellow or
mustard-yellow, 3–8 mm broad, 0.3–0.8 mm thick; surface of the
stromata smooth; perithecia completely immersed in the stroma,
obovate-subglobose, generally crowded and elongate, 150–200 μm high;
asci 48–56 × 2.6–3.0 μm; part-spores colourless, minutely
pustulate-tuberculate; generally globose to subglobose, often with a depressed
surface between paired part-spores, or the proximal part-spores often obovate,
sometimes ellipsoid, 2.2–3.2 × 2.0–2.8 μm
(<xref ref-type="bibr" rid="ref10">Doi 2001</xref>).</p><p><italic>Anamorph</italic>: Conidiophores verticillate, phialides in whorls of
3–5, slender, attenuated toward the tip, 14–32 ×
1.7–2.8 μm; conidia colourless, ovate, obovate, ellipsoid,
subcylindrical with a minutely truncate base, smooth, 2.6–6.2 ×
1.4–2.3 μm; chlamydospores not formed
(<xref ref-type="bibr" rid="ref10">Doi 2001</xref>).</p><p><italic>Habitat</italic>: Dead branches and twigs of <italic>Castanopsis
cuspidata</italic>.</p><p><italic>Known distribution</italic>: Japan.</p><p><italic>Specimens examined</italic>: Type material was not available for
examination.</p><p><italic>Comments</italic>: This species can be identified from the others discussed
here by its unique colour and pulvinate stroma. Pustulate-turberculate or
nodulose ascospores are not common among species of <italic>Hypocrea</italic> with
extensively effuse stromata and anamorphs assignable to <italic>Trichoderma</italic>sect. <italic>Hypocreanum</italic>. The minutely pustulate-tuberculate ascospores, and
elongate perithecia of <italic>H. mikurajimensis</italic> suggest a close relationship
to <italic>H. aurantiistroma, H. fulva</italic>, and <italic>H. megalocitrina</italic>.</p></list-item><list-item><p><bold><italic>Hypocrea pseudostraminea</italic></bold> Yoshim. Doi, Bull. Natl. Sci. Mus.
15: 676. 1972. Figs <xref rid="fig22" ref-type="fig">22</xref>,
<xref rid="fig23" ref-type="fig">23</xref>.</p><p><italic>Anamorph</italic>: <italic>Trichoderma</italic> sp. [sect.
<italic>Hypocreanum</italic>].</p><p><italic>Teleomorph</italic>: Stromata effuse, varying from solitary and discrete to
extensive confluent; largest continuous stroma 8 × 5 mm, smallest
continuous stroma 3 × 2 mm, with a white byssoid margin, varying in
colour, usually brownish orange to light brown (5C7–5E6);
KOH<sup>+/–</sup>, reaction variable, weak when hyphal protrusions not
abundant; KOH<sup>+</sup>, reaction strong, when hyphal protrusions abundant,
stroma typically turning reddish brown; ostiolar canals visible at stroma
surface, appearing brown. Stroma surface appearing granular due to minutely
roughened hyphal protrusions at the stroma surface; tissue immediately below
the stroma surface formed of loose pseudoparenchymatous cells and <italic>t.
intricata,</italic> anastomosing to form a more compact layer of <italic>t.
angularis</italic>; stroma tissue more angular near the centre of the stroma,
tissue near the margin becoming more hyphal. Pseudoparenchymatous tissue below
the perithecia formed of a loose <italic>t. globulosa</italic> and <italic>t.
intricata</italic>. Perithecia completely immersed, tightly compacted near the
stroma centre, fewer near the margin. Perithecia subglobose to ellipsoidal,
(133–)145–190(–207) μm long (including the length of the
ostiolar canal (n = 29); width of the perithecia near the base (measured from
3/4 total length of the perithecium), (74–)89–127 (–147)
μm (n = 29); length of ostiolar canal (32–)40–58 (–67)
μm; width of ostiolar canal (25–)29–41(–43) μm (n =
29); wall KOH<sup>+/–</sup>, reaction variable, weak, usually orangish
brown. Asci cylindrical, (48–)58–79(–92) ×
(3.3–)4.1–5.7(–7) μm (n = 158), tip slightly thickened.
Part-ascospores hyaline, spinulose, dimorphic; distal part subglobose,
sometimes subellipsoidal, (2.3–)3–4 (–5.5) ×
(2.3–)2.8–3.6(–4.2) μm, L/W ratio
(0.8–)0.97–1.2(–1.5) (n = 192); proximal part ellipsoidal,
sometimes subglobose to ovate, (2.2–)3.5–4.5(–6.2) ×
(2–)2.6–3.3(–4) μm, L/W ratio
(0.9–)1.2–1.6(–2) (n = 192).</p><p><italic>Anamorph</italic>: Colonies growing across the surface of a 100 mm diam
Petri plate with PDA in 10 d, with a dense white layer of mycelium close to
the agar surface and a uniform layer of cottony aerial mycelium above;
conidiophores visible in the aerial mycelium, borne on long elements of aerial
hyphae, irregularly branched; not all isolates producing conidiophores on PDA;
no odour; light brown pigment (7D7) near the edge of the plate; stroma
initials observed on PDA at 10 d in the same isolates that produce
conidiophores; chlamydospores not observed at 10 d. Colonies growing across
the surface of a 100 mm diam Petri plate with SNA in 10 d, producing a thin
layer of effuse mycelium covering the agar surface; no odour; no pigment; no
conidiophore production. Colonies growing across the surface of a 100 mm diam
Petri plate with CMD in 10 d, producing a thin layer of mycelium close to the
agar surface; conidiophores visible in the aerial mycelium, especially near
the edge of the plate; conidiophores verticillium-like, irregularly branched,
borne on long hyphal elements; no odour; no pigment production; stroma
initials (aggregates of mycelium leading to the production of stromata)
present; some isolates do not form conidiophores in culture;
chlamydospore-like swellings observed on CMD. Optimal temperature for growth
is 25–30 °C.</p><p>Measurements of micro-morphological characters were the same on all three
media. A description based on measurements from all three media at 10 d
follows: conidiophores irregularly verticillate, phialides solitary and
alternating, or in whorls of 3–5, subulate,
(9–)14–21(–36) × (1.5–)2.3–3.2(–3.8)
μm (n = 67); conidia variable, subellipsoidal to elongate ellipsoidal,
(3–)4.7–6.4(–10) × (2–)2.4–3.1(–3.4)
μm (n = 67), sometimes with asymmetric, truncate base.</p><p><italic>Habitat</italic>: On bark of <italic>Quercus, Rhododendron</italic>, possibly
lichenicolous, possibly fungicolous as found growing on <italic>Armillaria</italic>rhizomorphs.</p><p><italic>Known distribution</italic>: Japan, North America, and France.</p><p><italic>Paratypes examined</italic>: <bold>Japan,</bold> Hirakura, the Forestry
Experiment Station, Mie Pef., on bark, 7 Sep. 1965, Yoshim. Doi (NY ex
TNS.D-56 = TNSF 19051); Tsushima Island, Nagasaki Pref., on bark, 27 Sep.
1968, Yoshim. Doi (NY ex TNS.D-487 = TNSF 223484).</p><p><italic>Other specimens examined</italic>: <bold>France,</bold> Osserain, on
<italic>Phyllostachys</italic> sp., 22 Oct. 1989, F. Candoussau No. 4805-16 (BPI
1107143). <bold>U.S.A.,</bold> Indiana, Brown Co., vic. of Pike's Peak, Happy Hollow
Camp, 39°09' N, 86°06' W, elev. 250 m, on root, 29 Sep. 1995, G. J.
Samuels (BPI 737763; culture G.J.S. 95-189); Kentucky, Daniel Boone National
Forest, Cave Run Lake, Sheltowee Trail from Stone Cave, on decayed lichen on
bark, 26 Sep. 1995, G. J. Samuels (BPI 737729, culture G.J.S. 95-169);
Maryland, Howard Co. and Carroll Co. border, Paptapsco State Park, outside of
Sykesville, on rotten wood and <italic>Armillaria</italic> sp. rhizomorphs, 12 Oct.
1999, B.E. Overton, B.E.O. 99-36 (BPI; culture B.E.O. 99-36); Prince Georges
Co., E. of Largo in old growth forest at Church Rd., on decorticated wood, 11
Oct. 1991, G. J. Samuels, S. Rehner, A.Y. Rossman, & F.A. Uecker (BPI
1112904); New York, Dutchess Co., E. side of Pawling Nature Reserve, on bark,
6 Oct. 1990, G.J. Samuels & C.T. Rogerson (BPI 1107187; culture G.J.S.
90-74).</p><p><italic>Comments</italic>: Specimen B.E.O. 99-36 has roughened hyphal protrusions
near the stroma surface that are more prevalent and developed than in the
other specimens examined and it was the only collection from those examined on
<italic>Armillaria</italic> sp. rhizomorphs. In addition, molecular data suggest that
this collection represents a unique phylogenetic species. However, the
ascospore measurements and anamorph characteristics of B.E.O. 99-36 are
identical to the type of <italic>H. pseudostraminea.</italic> At this time, it is
unclear whether this collection represents a unique species as more isolates
have to be sequenced. The orange-brown stromata, granular appearance of the
stromata, and part-ascospore measurements allow <italic>H. pseudostraminea</italic> to
be distinguished <italic>from H. microcitrina. Hypocrea lactea sensu</italic> Doi has
hyphal protrusions near the stroma surface similar to those observed in <italic>H.
pseudostraminea</italic>.</p></list-item><list-item><p><bold><italic>Hypocrea microcitrina</italic></bold> Yoshim. Doi, Bull. Natl. Sci. Mus.
15: 667. 1972. Figs <xref rid="fig24" ref-type="fig">24</xref>,
<xref rid="fig25" ref-type="fig">25</xref>.</p><p><italic>Anamorph</italic>: <italic>Trichoderma</italic> sp. [sect.
<italic>Hypocreanum</italic>].</p><p><italic>Teleomorph</italic>: Stromata effuse, extensive and confluent, largest
continuous stroma 12 × 11 mm, smallest continuous stroma 2 × 2 mm,
with a white byssoid margin, varying in colour, greyish yellow to orangish
brown, sometimes light yellow (4B5–5E4, 3A4), ostiolar canals visible at
stroma surface, appearing orange (5A4). KOH<sup>+</sup>, reaction variable,
stromata becoming darker orange. Stroma surface appearing granular due to
minutely roughened hyphal protrusions, tissue at stroma surface formed of
loose to compact <italic>t. intricata</italic>. Pseudoparenchymatous tissue below the
perithecia of <italic>t. intricata</italic> or <italic>t. globulosa</italic>, variable among
collections, sometimes compact or sometimes in loose aggregates. Perithecia
completely immersed, tightly compacted near the stroma centre, fewer near the
margin. Perithecia subglobose to ellipsoidal,
(176–)180–205(–210) 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); (80–)88–126(–140) μm (n
= 16); length of ostiolar canal (33–)38–53(–58) μm; width
of ostiolar canal (35–)38–50(–53) μm (n = 16); wall
KOH<sup>+/–</sup>, reaction variable, usually becoming dark orange. Asci
cylindrical, (48–)53–67(–77) × (3.3–)
3.9–4.9(–5.8) μm (n = 105), tip slightly thickened.
Part-ascospores hyaline, thick-walled, spinulose, dimorphic; distal part
subglobose, sometimes subellipsoidal, (2.3–) 2.7–3.5(–4)
× (2.3–)2.7–3.3(–3.8) μm, L/W ratio
(0.75–)0.9–1.2(–1.4) (n = 140); proximal part, ellipsoidal,
sometimes subglobose to ovoidal, (2.6–)3.1–3.9(–4.7) ×
(1.9–)2.4–2.9(–3.6) μm, L/W ratio
(0.9–)1.1–1.5(–1.8) (n = 192).</p><p><italic>Anamorph</italic>: Colonies growing across the surface of a 100 mm diam
Petri plate with PDA in 10 d; not producing concentric rings or radial rays of
mycelium; a dense layer of cottony aerial mycelium covering the entire Petri
plate; sometimes aerial mycelium producing yellowish brown exudates;
conidiophores not visible in the aerial mycelium at 10 d; only some
conidiophore aggregates near the inoculation plug and the edge of the plate;
conidiophores irregularly verticillate; phialides subulate, typically in
whorls of 3–5, but also solitary or in groups of two,
(8–)13–22(–33) × (1.3–)2.3–3.2(–4.3)
μm (n = 50); conidia hyaline, smooth, ellipsoidal to elongate ellipsoidal,
variable in size, (3.7–)4.8–6(–12) ×
(1.9–)2.3–2.9(–3.6) μm (n = 69), without a well-defined
flat edge, rarely with a septum; chlamydospores not observed at 10 d on PDA;
culture producing yellowish grey diffusible pigment (4B6), near the agar plug
appearing darker orange (5B8) to brown (5D7). After 10 d conidia begin to
swell, becoming larger and more variable in size. Colonies growing across the
surface of a 100 mm diam Petri plate with SNA in 10 d; a thin layer of effuse
mycelium covering the whole surface; no odour, no pigment production; no
conidiophore production. Colonies growing across the surface of a 100 mm diam
Petri plate with CMD in 10 d, producing a thin layer of effuse mycelium
covering the agar; no conidiophore production, no odour, and no pigment
production. Optimal temperature for growth is 25 °C on all three
media.</p><p><italic>Habitat</italic>: On bark, possibly fungicolous.</p><p><italic>Known distribution</italic>: Japan and North America.</p><p><italic>Type</italic>: The type of <italic>H. microcitrina</italic> was not available for
study.</p><p><italic>Specimens examined</italic>: <bold>U.S.A.,</bold> Georgia, Rayburn Co.,
Chatachoochie National Forest, vic. Clayton, Warwomen Dell., elev. 750 m, E.
Lieckfeldt & G. J. Samuels (BPI 744660; culture G.J.S. 97-248); New
Jersey, Newfield, on leaf, Oct. 1894, Ellis & Everhart 3217 (BPI 631561);
Virginia, Giles Co., Mt. Lake Biological Station, vic. Little Spruce Bog,
37°22' N, 80°31' W, elev. 1170 m, growing on black perithecial fungus
on bark, 17 Sep. 1991, G. J. Samuels, C. Rogerson, S. Huhndorf, S. Rehner
& M. Williams (BPI 1112833; culture G.J.S. 91-61); Giles Co., Mt. Lake
Biological Station, Horse Barn Trail, 37°22' N, 80°3' W, on
decorticated wood, 19 Sep. 1991, G. J. Samuels, C. Rogerson, S. Huhndorf, S.
Rehner & M. Williams (BPI 1112839).</p><p><italic>Comments</italic>: <italic>Hypocrea microcitrina</italic> and <italic>H.
pseudostraminea</italic> are similar species and difficult to distinguish
(<xref ref-type="bibr" rid="ref9">Doi 1972</xref>). The stroma surface
of <italic>H. microcitrina</italic> is composed of <italic>t. intricata</italic> with hyphae
that do not anastomose to form a thick-walled <italic>t. globulosa</italic> or <italic>t.
angularis</italic> as in <italic>H. pseudostraminea</italic>. It is difficult to interpret
whether the hyphal elements near the stroma surface of <italic>H.
microcitrina</italic> are hyphal protuberances as in <italic>H. pseudostraminea</italic>or just a loose layer of <italic>t. intricata</italic>. Regardless of homology, the
hyphal elements in <italic>H. microcitrina</italic> macroscopically make the stroma
surface appear granular as in <italic>H. pseudostraminea. Hypocrea
microcitrina</italic> has slightly smaller ascospores than <italic>H. pseudostraminea.
Hypocrea microcitrina</italic> was previously only known from Japan and has not
previously been reported from the United States, where it also appears to be
common on woody substrata.</p></list-item><list-item><p><bold><italic>Hypocrea albocitrina</italic></bold> Yoshim. Doi, Bull. Natl. Sci. Mus. 15:
661. 1972. <xref rid="fig26" ref-type="fig">Fig. 26</xref>.</p><p><italic>Anamorph</italic>: <italic>Trichoderma</italic> sp. [sect.
<italic>Hypocreanum</italic>].</p><p><italic>Teleomorph</italic>: Stromata effuse, solitary to confluent, largest
continuous stromata 5 × 3 mm, smallest continuous stromata 1 × 1
mm, stromata pastel-yellow (3A4), KOH<sup>–</sup>, tissue below the
stroma surface of <italic>t. globulosa</italic> to <italic>t. intricata</italic>, loosely
arranged; ostiolar canals visible from the stroma surface, appearing yellow;
perithecia globose, widely spaced, not abundant. Asci cylindrical,
(63–)66–78(–83) × (4.8–)4.9–5.7(–6)
μm (n = 17), tip slightly thickened. Asci with (12–14–)16
part-ascospores; ascospores often segregating into 4 large or 4 small spores;
segregation pattern of part<bold>-</bold>spores 4 large to 4 small and 8 large to 8
small common, with some 4 to 8 to 4. Asci with fewer than 16 part-spores
usually showing segregation patterns of 2 to 8 to 2, or 2 to 8 to 4; the
smaller part-ascospores then appearing to be aborted or missing; regardless of
segregation pattern, the smaller part-ascospores are not as mature or
well-formed as the larger part-ascospores. Part-ascospores hyaline,
thick-walled, spinulose; smaller part-spores subglobose to subellipsoidal,
(2.3–)2.8–3.6(–3.8) ×
(2.4–)3–3.7(–4.0) μm, L/W ratio
(0.7–)0.84–1.08(–1.4) (n = 38); larger part-spores
subglobose to ellipsoidal, (3.9–)4.3–5.1(–5.5) ×
(3.3–)3.7–4.5(–4.8) μm, L/W ratio
(0.9–)1–1.3(–1.5) (n = 38).</p><p><italic>Anamorph</italic>: verticillium-like (fide
<xref ref-type="bibr" rid="ref9">Doi 1972</xref>). Conidiophores
irregularly verticillate, forming minute colonies; phialides slender,
13–45 × 2–4 μm; conidia hyaline, smooth, cylindrical to
ellipsoidal, with a truncate base, 2.8–19 × 2–4 μm
(<xref ref-type="bibr" rid="ref9">Doi 1972</xref>).</p><p><italic>Habitat</italic>: On decayed branches of <italic>Abies firma</italic>.</p><p><italic>Known distribution</italic>: Japan.</p><p><italic>Isotype</italic>: <bold>Japan,</bold> Tsushima Island, Nagasaki Pref., Mt.
Mitake, on bark, 28 Sep. 1968, Y. Doi (NY ex TNS-F-190517).</p><p><italic>Comments</italic>: The isotype material is in good condition. The
segregation of ascospores based on size is a unique character that
distinguishes this species from others in subsect. <italic>Citrinae</italic>.</p></list-item><list-item><p><bold><italic>Hypocrea protocitrina</italic></bold> Yoshim. Doi, Bull. Natl. Sci. Mus.
15: 660. 1972.</p><p><italic>Anamorph</italic>: Unknown.</p><p><italic>Teleomorph</italic>: Stromata effuse, thin, yellow, up to 4 × 12 mm,
200–240 μm thick; internal tissue of the stromata of <italic>t.
intricata</italic>; perithecia entirely embedded in the stroma,
obovate-subglobose, 110–130 μm high; perithecial necks, 20–30
μm long; part-ascospores hyaline, very minutely warted; distal part
subglobose to obovate-ellipsoid, 1.9–2.5 × 1.7–2.1 μm;
proximal part obovate-ellipsoid, 2.5–3.5 × 1.6–2.0 μm
(<xref ref-type="bibr" rid="ref9">Doi 1972</xref>).</p><p><italic>Habitat</italic>: On decayed bamboo.</p><p><italic>Known distribution</italic>: Japan.</p><p><italic>Isotype</italic>: <bold>Japan,</bold> Tsushima Island, Nagasaki Pref., Toshibachi
Izuhara-Cho, on bamboo, 26 Sep. 1968, Y. Doi (NY ex TNS D-482).</p><p><italic>Comments</italic>: The isotype material is immature and limited in quantity
not allowing for destructive sampling. The minute ascospores of <italic>H.
protocitrina</italic> distinguish this species from all others that Doi
(<xref ref-type="bibr" rid="ref9">1972</xref>) included in subsect.
<italic>Citrinae</italic>.</p></list-item></list></p><p><fig position="float" id="fig5"><label>Fig. 5.</label><caption><p>A–F. <italic>Hypocrea citrina</italic>. A. Macromorphology of B.E.O. 99-29;
bar = 2 mm. B. Stromatal section showing <italic>t. globulosa</italic> to <italic>t.
angularis</italic> tissues intermixed with <italic>t. intricata</italic> in BPI 737759;
bar = 20 μm. C. Stroma variable in colour, white to yellow,
<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> isolated
from specimen (photo courtesy of W. Gams). D. Asci with ascospores of neotype
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=4031&link_type=CBS">CBS 4031</ext-link>; bar = 20
μm. E. Stromatal cross section showing <italic>t. globulosa</italic> with angular
edges due to mutual compaction in BPI 748251; bar = 20 μm. F. Immature asci
and ascospores of BPI 748251; bar = 20 μm.</p></caption><graphic xlink:href="1fig5"></graphic></fig></p><p><fig position="float" id="fig6"><label>Fig. 6.</label><caption><p>A–C. <italic>Hypocrea citrina</italic>. A. Neotype specimen Gams 4031,
<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> on leaf
litter. B. Stromatal section of BPI 744475 showing <italic>textura angularis</italic>below the perithecium and near the stroma surface; bar = 40 μm. C. Variable
KOH reaction usually not greater than observed here in BPI 744475; bar = 2
mm.</p></caption><graphic xlink:href="1fig6"></graphic></fig></p><p><fig position="float" id="fig7"><label>Fig. 7.</label><caption><p>A–K. <italic>Hypocrea citrina</italic> (specimens of <italic>Hypocrea
lactea</italic>). A. Illustration of <italic>Sphaeria lactea</italic> from Fries, Obs.
Taf. 8, fig. 4. Compare to B. B–G. <italic>Sphaeria lactea,</italic> neotype
specimen, coll. E.M. Fries, UPS; B. specimen glued to a piece of stiff paper.
C. Surface of stroma seen in face view. D. Macroscopic view of stroma surface;
bar = 1 mm. E. Ascus with developing ascospores; bar = 20 μm. F. Immature
asci showing pore in ascus apex (left, cotton blue in lactic acid, right in 1
% aq. phloxine); bar = 10 μm. G. Mature ascospores in asci (cotton blue in
lactic acid); bar = 10 μm. H–K. <italic>Hypocrea lactea</italic> specimen
collected by Fries, Femsjö, UPS. H. Specimen; bar = 1 cm. I. Perithecial
papilla, ostiolar opening and stroma surface; bar = 20 μm. J. Stroma
surface showing pseudoparenchymatous cells at the surface and loosely disposed
hyphae below; bar = 20 μm. K. Mature ascospores in asci (cotton blue in
lactic acid); bar = 10 μm.</p></caption><graphic xlink:href="1fig7"></graphic></fig></p><p><fig position="float" id="fig8"><label>Fig. 8.</label><caption><p>A–G. Anamorph of <italic>Hypocrea citrina.</italic> A. Aerial mycelium with
irregular verticillium-like conidiophores,
<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> on PDA;
bar = 20 μm. B, D. Bulbous terminal and globose intercalary chlamydospores,
B.E.O. 99-29 on PDA; bar = 20 μm. C, G. Variation in conidial size and
shape, B.E.O. 99-29 on PDA and
<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> on PDA,
respectively; bar = 20 μm. E–F. Variation in phialide arrangement
with typically 3–5 phialides per whorl,
<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> on PDA
and B.E.O. 99-29, respectively; bar = 20 μm.</p></caption><graphic xlink:href="1fig8"></graphic></fig></p><p><fig position="float" id="fig9"><label>Fig. 9.</label><caption><p>A–C. <italic>Hypocrea pulvinata</italic>, BPI 744722. A. Subpulvinate
stromata; bar = 2 mm. B. Orange brown subpulvinate stromata and positive KOH
reaction; bar = 2 mm. C. Ostiolar openings visible at stroma surface; bar = 1
mm.</p></caption><graphic xlink:href="1fig9"></graphic></fig></p><p><fig position="float" id="fig10"><label>Fig. 10.</label><caption><p>A–F. <italic>Hypocrea pulvinata</italic>, BPI 744722. A. Section through
stroma; bar = 40 μm. B. Ostiolar canal with pointed papillae; bar = 20
μm. C. Warted hyphal protuberances at stroma surface; bar = 20 μm. D.
Vertically elongate <italic>t. angularis</italic> to tightly packed <italic>t.
intricata</italic> below perithecia; bar = 20 μm. E–F. Asci with
ascospores; bar = 20 μm.</p></caption><graphic xlink:href="1fig10"></graphic></fig></p><p><fig position="float" id="fig11"><label>Fig. 11.</label><caption><p>A–C. Anamorph of <italic>Hypocrea pulvinata</italic>, G.J.S. 98-104. A.
Solitary phialides on long hyphal branches, on PDA; bar = 20 μm. B.
Chlamydospore-like swellings below phialides, on PDA; bar = 20 μm. C.
Multiple phialides on branched hyphal elements, on PDA; bar = 20 μm.</p></caption><graphic xlink:href="1fig11"></graphic></fig></p><p><fig position="float" id="fig12"><label>Fig. 12.</label><caption><p>A–C. <italic>Hypocrea americana</italic>, CUP 38045. A. Subpulvinate stroma;
bar = 2 mm. B. Stromata with rugulose appearance, ostiolar openings not
readily visible; bar = 1 mm. C. Colour reaction in KOH; bar = 1 mm.</p></caption><graphic xlink:href="1fig12"></graphic></fig></p><p><fig position="float" id="fig13"><label>Fig. 13.</label><caption><p>A–E. <italic>Hypocrea americana</italic>, CUP 38045. A. Section of stroma
showing tissue types; bar = 40 μm. B, E. Asci with ascospores; bar = 20
μm. C. Warted hyphal protuberances near stroma surface; bar = 20 μm. D.
Tissue below perithecia; bar = 20 μm.</p></caption><graphic xlink:href="1fig13"></graphic></fig></p><p><fig position="float" id="fig14"><label>Fig. 14.</label><caption><p>A–D. Anamorph of <italic>H. americana.</italic> A. Chlamydospore-like
swellings below phialides, on PDA. B. Phialides on long hyphal elements, on
SNA, both G.J.S. 96-191. C. Variability in conidial size and shape, on CMD,
G.J.S. 92-93. D. Conidiophores with multiple branches, on CMD, G.J.S. 96-191;
bars = 20 μm.</p></caption><graphic xlink:href="1fig14"></graphic></fig></p><p><fig position="float" id="fig15"><label>Fig. 15.</label><caption><p>A–G. <italic>Hypocrea protopulvinata.</italic> A. Effuse to subpulvinate
stromata, K.P. 00-56; bar = 2 mm. B. Reaction of stromata in KOH, B.E.O.
01-01. C. Asci with ascospores; bar = 20 μm. D. Roughened hyphal
protuberances near stroma surface and <italic>t. intricata</italic>, bar = 20 μm;
C. and D. TNS 223431. E. Cross section of perithecium; bar = 20 μm. F.
Tissue near ostiolar canals of <italic>t. globulosa</italic> approaching <italic>t.
angularis</italic>; bar = 20 μm; E. and F. K.P. 00-56. G. Tissue below base of
perithecium, <italic>t. globulosa</italic>, sometimes with angular edges due to mutual
compaction, TNS 223431; bar = 20 μm.</p></caption><graphic xlink:href="1fig15"></graphic></fig></p><p><fig position="float" id="fig16"><label>Fig. 16.</label><caption><p>A–F. Anamorph of <italic>H. protopulvinata.</italic> A. Conidiophores arising
from compact hyphal aggregates, on PDA; bar = 20 μm. B. Conidiophores
arising from long hyphal elements, on CMD; bar = 20 μm. C. Branched
conidiophores, on CMD; bar = 20 μm; A–C.
<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>.
D–E. Variation of conidial size and shape in K.P. 00-56 on PDA, and
<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> on CMD,
respectively; bar = 20 μm. F. Branched conidiophores with slight swelling
of hyphae below phialides, K.P. 00-56, on PDA; bar = 20 μm.</p></caption><graphic xlink:href="1fig16"></graphic></fig></p><p><fig position="float" id="fig17"><label>Fig. 17.</label><caption><p>A–E. <italic>Hypocrea megalocitrina.</italic> A. Effuse stromata with
projecting ostiolar canals, B.E.O. 99-42; bar = 2 mm. B. Stroma
characteristics of the type, TNS 223220; bar = 2 mm. C. Variation in conidial
size and shape; bar = 20 μm. D. Verticillium-like anamorph; bar = 20 μm.
E. Phialides in whorls, usually of 3–5; bar = 20 μm; C–E.
B.E.O. 00-09.</p></caption><graphic xlink:href="1fig17"></graphic></fig></p><p><fig position="float" id="fig18"><label>Fig. 18.</label><caption><p>A–F. <italic>Hypocrea megalocitrina.</italic> A. Section of stroma showing
protruding ostiolar canals and tissue types, B.E.O. 99-42; bar = 40 μm. B.
Discharged nodulose ascospores, TNS 223220; bar = 20 μm. C. Tightly packed
<italic>t. intricata</italic> near stroma surface, sometimes appearing angular when
tightly packed, sometimes loosely woven; bar = 20 μm. D. Vertically
elongate <italic>t. globulosa</italic> to <italic>t. intricata</italic> below perithecium, no
space between the hyphae; bar = 20 μm; C–D. L.G.1. E, F. Asci and
conical ascospores, TNS 223220 and L.G.1, respectively; bar = 20 μm.</p></caption><graphic xlink:href="1fig18"></graphic></fig></p><p><fig position="float" id="fig19"><label>Fig. 19.</label><caption><p>A–D. <italic>Hypocrea aurantiistroma</italic>, BPI 1107193. A. Ostiolar
canals projecting from stroma surface; bar = 1 mm. B. Stroma surface including
margin; bar = 2 mm. C–D. Asci and minutely nodulose ascospores; bar = 20
μm.</p></caption><graphic xlink:href="1fig19"></graphic></fig></p><p><fig position="float" id="fig20"><label>Fig. 20.</label><caption><p>A–D. <italic>Hypocrea aurantiistroma</italic>, BPI 1107193. A. Section
through stroma; bar = 40 μm. B. <italic>Textura intricata</italic> near ostiolar
canal; bar = 20 μm. C. Cross section of protruding ostiolar canal; bar = 20
μm. D. Loose <italic>t. globulosa</italic> to tightly packed <italic>t. intricata</italic>below perithecium; bar = 20 μm.</p></caption><graphic xlink:href="1fig20"></graphic></fig></p><p><fig position="float" id="fig21"><label>Fig. 21.</label><caption><p>A–E. <italic>Hypocrea fulva</italic>, syntype 410. A. Stromata; bar = 2 mm.
B. Section of stroma surface including margin; bar = 40 μm. C–D.
Cross sections showing stroma anatomy; bar = 40 μm. E. Asci and minutely
nodulose ascospores; bar = 20 μm.</p></caption><graphic xlink:href="1fig21"></graphic></fig></p><p><fig position="float" id="fig22"><label>Fig. 22.</label><caption><p>A–E. <italic>Hypocrea pseudostraminea.</italic> A. Effuse to subpulvinate
stroma with white byssoid margin; bar = 1 mm. B. Reaction in KOH; bar = 1 mm;
both TNS 223484. C. KOH-positive roughened hyphal protuberances, B.E.O. 99-36;
bar = 20 μm. D. Asci and ascospores, TNS 223484; bar = 20 ìm. E.
Cross section showing tissue near ostiolar canal and below perithecium, BPI
1112904; bar = 40 μm.</p></caption><graphic xlink:href="1fig22"></graphic></fig></p><p><fig position="float" id="fig23"><label>Fig. 23.</label><caption><p>A–C. Anamorph of <italic>H. pseudostraminea</italic>, G.J.S. 91-135. A.
Verticillium-like anamorph, on CMD. B. Variation in conidial size and shape,
on CMD. C. Conidiophores on long hyphal element, on CMD; bars = 20 μm.</p></caption><graphic xlink:href="1fig23"></graphic></fig></p><p><fig position="float" id="fig24"><label>Fig. 24.</label><caption><p>A–C. <italic>Hypocrea microcitrina</italic>, BPI 1112833. A. Effuse stroma
with white byssoid margin; bar = 2 mm. B. Section of stroma in KOH; bar = 40
μm. C. KOH reaction of stroma; bar = 1 mm. D–E. Section showing
tissue types; bar = 40 μm. F. Tissue below perithecium composed of <italic>t.
globulosa</italic> with angular edges due to mutual compaction; bar = 20 μm. G.
Ascus with ascospores; bar = 20 μm.</p></caption><graphic xlink:href="1fig24"></graphic></fig></p><p><fig position="float" id="fig25"><label>Fig. 25.</label><caption><p>A–F. Anamorph of <italic>H. microcitrina.</italic> A. Sterile aerial
mycelium, G.J.S. 91-61; bar = 20 μm. B. Irregular verticillium-like
anamorph with variable branching pattern, on PDA; bar = 20 μm. C–D.
Phialides in varying numbers, usually not more than 3 per node, on PDA; bar =
20 μm. E. Variation in conidial size and shape, on PDA; bar = 20 μm. F.
Solitary phialides on long hyphal extensions, on PDA; bar = 20 μm;
B–F. G.J.S. 97-248.</p></caption><graphic xlink:href="1fig25"></graphic></fig></p><p><fig position="float" id="fig26"><label>Fig. 26.</label><caption><p>A–F. <italic>Hypocrea albocitrina</italic>, TNS 190517. A. Effuse stromata;
bar = 1 mm. B. Ostiolar canals visible at stroma surface; bar = 1.5 mm. C.
Section of stroma showing tissue types; bar = 40 μm. D. Asci with
ascospores segregating, different sizes; bar = 20 μm. E–F. Stroma
tissue composed of loosely woven <italic>t. globulosa</italic> and <italic>t.
intricata</italic> near the stroma surface and below the perithecium; bar = 20
μm.</p></caption><graphic xlink:href="1fig26"></graphic></fig></p></sec><sec><title>KEY TO <italic>HYPOCREA CITRINA</italic> AND ALLIES</title><p><list list-type="simple"><list-item><p>1. Stromata attached by a narrow base; part-ascospores spinulose,
monomorphic, globose, 2.1–2.3 μm diam, or sometimes
obovate-ellipsoid, 2.0–3.0 μm long...................................
5. <bold><italic>H. platypulvinata</italic></bold></p></list-item><list-item><p>1. Stromata effuse to
pulvinate...............................................................................................................................
2</p></list-item><list-item><p>2. Part-ascospores monomorphic without ornamentation; on
polypores................................................................
3</p></list-item><list-item><p>2. Part-ascospores not as above; mostly on other
substrata.................................................................................
4</p></list-item><list-item><p>3. Part-ascospores ellipsoidal, (2.8–)3.8–5(–6.7) ×
(2.3–)2.8-3.8(–4.7) μm..................................... 2.
<bold><italic>H. pulvinata</italic></bold></p></list-item><list-item><p>3. Part-ascospores, subglobose to rhomboidal,
(1.7–)2.6–3.8(–6.2) ×
(2.4–)3.2–4.2(–5.4)
μm............................................................................................................
3. <bold><italic>H. americana</italic></bold></p></list-item><list-item><p>4. Part-ascospores smooth, dimorphic; distal part subglobose,
(3–)3.3–3.9 (–4.3) ×
(2.6–)3–3.5(–4.2) μm; proximal part ellipsoidal,
sometimes subglobose, (3.2–)3.0–4.2 (–4.8) ×
(2.4–)2.7–3.3(–3.8) μm; on polypores (<italic>Fomitopsis</italic>and others).............................. 5. <bold><italic>H. protopulvinata</italic></bold>[If part-ascospores slightly smaller and anamorph <italic>Gliocladium</italic>; on
soft polypores (e.g. <italic>Tyromeces</italic>), see <italic>H. pallida</italic>]</p></list-item><list-item><p>4. Part-ascospores spinulose or nodulose; not on
polypores.................................................................................
5</p></list-item><list-item><p>5. Stromata white to yellow, effuse, and with pseudoparenchymatous tissue
near the surface; on leaf litter, soil, bark, or moss; part-ascospores
spinulose, distal part subglobose, sometimes subellipsoidal,
(3.2–)4.1–5.2(–6.3) ×
(2.9–)3.7–4.8(–5.7) μm; proximal part ellipsoidal,
sometimes subglobose to ovate, (3.3–)4.4–6(–7.1) ×
(2.5–)3.3–4.9(–6.3) μm................1. <bold><italic>H.
citrina</italic></bold></p><p>5. Stromata variable in colour, effuse or pulvinate, hyphal to
pseudoparenchymatous; growing on decorticated wood, or old fructifications of
fungi on bark; part-ascospores spinulose or
nodulose........................................................................................................................................
6</p></list-item><list-item><p>6. Stroma surface with hyphal protuberances, extensive in some specimens,
minimal in others, arising from a well-defined layer of pseudoparenchymatous
tissue, KOH<sup>+/–</sup>, variable, stromata orange-brown to light
brown; on bark, possibly fungicolous; part-ascospores spinulose; distal part
subglobose, sometimes subellipsoidal, (2.3–)3–4(–5.5)
× (2.3–)2.8–3.6(–4.2) μm; proximal part
ellipsoidal, sometimes subglobose to ovate,
(2.2–)3.5–4.5(–6.2) ×
(2–)2.6–3.3(–4) μm.................................. 10.
<bold><italic>H. pseudostraminea</italic></bold></p></list-item><list-item><p>6. Stroma surface without hyphal protuberances arising from a
pseudoparenchymatous layer near the
surface............................................................................................................................
7</p></list-item><list-item><p>7. Part-ascospores
spinulose.................................................................................................................................
8</p></list-item><list-item><p>7. Part-ascospores
nodulose.................................................................................................................................
10</p></list-item><list-item><p>8. On bark; stroma surface composed of <italic>t. intricata</italic>, greyish
yellow to orange-brown; distal part-ascospores subglobose, sometimes
subellipsoidal, (2.3–)2.7–3.5(–4) ×
(2.3–)2.7–3.3(–3.8) μm; proximal part ellipsoidal,
sometimes subglobose to ovoidal, (2.6–)3.1–3.9(–4.7) ×
(1.9–)2.4–2.9(–3.6)
μm......................................................... 11. <bold><italic>H.
microcitrina</italic></bold></p></list-item><list-item><p>8. On bamboo or with eight large and eight small
part-ascospores........................................................................
9</p></list-item><list-item><p>9. On bamboo; stroma surface composed of <italic>t. intricata</italic>, yellow;
part-ascospores hyaline, minutely warted; distal part subglobose to
obovate-ellipsoid, 1.9–2.5 × 1.7–2.1 μm, proximal part
obovate-ellipsoid, 2.5–3.5 × 1.6–2.0
μm................................... 13. <bold><italic>H.
protocitrina</italic></bold></p></list-item><list-item><p>9. On decayed tree branches; stroma surface of loose to compated <italic>t.
globulosa</italic> with <italic>t. intricata</italic>, yellow; with eight large and eight
small, hyaline part-ascospores; smaller part-spores subglobose to
subellipsoidal, (2.3–)2.8–3.6(–3.8) ×
(2.4–)3–3.7(–4.0) μm; larger part-spores subglobose to
ellipsoidal, (3.9–)4.3–5.1(–5.5) ×
(3.3–)3.7–4.5(–4.8)
μm............................................................................................................................
12. <bold><italic>H. albocitrina</italic></bold></p></list-item><list-item><p>10. Stroma surface effuse to subpulvinate, composed mostly of <italic>t.
intricata</italic> with ostiolar canals projecting from stroma
surface..............................................................................................................
11</p></list-item><list-item><p>10. Stroma surface pulvinate with pseudoparenchymatous
tissue.........................................................................
12</p></list-item><list-item><p>11. Stroma surface greyish orange; part-ascospores slightly dimorphic,
almost appearing monomorphic; distal part subellipsoidal, slightly conical,
(3.6–)4.5–5(–5.9) ×
(2.5–)3.3–4(–4.6) μm; proximal part ellipsoidal, rarely
subellipsoidal, slightly conical, (3.6–)4.2–5.4(–6.4)
× (3.1–)3.3-4(–4.6)
μm.........................................................................
7. <bold><italic>H. aurantiistroma</italic></bold></p></list-item><list-item><p>11. Stroma surface light brown; part-ascospores dimorphic, distal part
subellipsoidal to conical, (3.2–)4.5–5.8(–6.6) ×
(3–)3.8–5(–6) μm; proximal part ellipsoidal, rarely
subellipsoidal, sometimes appearing thimble-shaped or conical,
(3.7–)4.6–5.9(–7) ×
(3.1–)3.7–4.8(–5.8)
μm.........................................................................
6. <bold><italic>H. megalocitrina</italic></bold></p></list-item><list-item><p>12. Part-ascospores dimorphic; distal part subellipsoidal, slightly
conical, (3.9–)4.5–5.5(–6.2) ×
(2.8–)3.2–3.8(–4.3) μm; proximal part ellipsoidal, rarely
conical, (4.3–)4.6–5.6(–6.5) ×
(2.8–)3.0–3.6 (–4.1); stromata
yellow-brown....................................................................................
8. <bold><italic>H. fulva</italic></bold></p></list-item><list-item><p>12. Part-ascospores monomorphic, globose to subglobose, 2.2–3.2
× 2.0–2.8 μm; stromata
mustard-yellow...................................................................................................
9. <bold><italic>H. mikurajimensis</italic></bold></p></list-item></list></p></sec></body><back><ack><p>We wish to acknowledge the following persons for their contributions to
this manuscript: Dr Gary J. Samuels for his comments on the systematics of
<italic>H. citrina</italic>, for the photographs of <italic>H. lactea</italic>, and access to
specimens, Dr Ronald Petersen for his assistance in questions of nomenclature,
Dr E. Parmasto for his identification of polypores in our study, Dr Elke
Lieckfeldt (deceased Aug. 2002) for her support and comments with the initial
planning stages of the molecular portion of this project, Dr Yoshimichi Doi
for making specimens available for study during the first author's work in
Japan, and Dr Walter Gams for rendering the Latin diagnoses and his editorial
comments on the manuscript. Dr. James Bray identified the moss
<italic>Drepanocladus</italic> <italic>uncinatus</italic> (Hedw.) Warnst. This study was
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