A molecular phylogenetic reappraisal of the Hysteriaceae, Mytilinidiaceae and Gloniaceae (Pleosporomycetidae, Dothideomycetes) with keys to world species
Identifieur interne : 000261 ( Pmc/Corpus ); précédent : 000260; suivant : 000262A molecular phylogenetic reappraisal of the Hysteriaceae, Mytilinidiaceae and Gloniaceae (Pleosporomycetidae, Dothideomycetes) with keys to world species
Auteurs : E. W. A. Boehm ; G. K. Mugambi ; A. N. Miller ; S. M. Huhndorf ; S. Marincowitz ; J. W. Spatafora ; C. L. SchochSource :
- Studies in Mycology [ 0166-0616 ] ; 2009.
Abstract
A reappraisal of the phylogenetic integrity of bitunicate ascomycete fungi
belonging to or previously affiliated with the
Url:
DOI: 10.3114/sim.2009.64.03
PubMed: 20169023
PubMed Central: 2816966
Links to Exploration step
PMC:2816966Le document en format XML
<record><TEI><teiHeader><fileDesc><titleStmt><title xml:lang="en">A molecular phylogenetic reappraisal of the <italic>Hysteriaceae</italic>,
<italic>Mytilinidiaceae</italic> and <italic>Gloniaceae</italic> (<italic>Pleosporomycetidae</italic>,
<italic>Dothideomycetes</italic>) with keys to world species</title><author><name sortKey="Boehm, E W A" sort="Boehm, E W A" uniqKey="Boehm E" first="E. W. A." last="Boehm">E. W. A. Boehm</name><affiliation><nlm:aff id="aff1"><italic>Department of Biological Sciences, Kean University, 1000 Morris Ave., Union, New Jersey 07083, U.S.A.</italic></nlm:aff></affiliation></author><author><name sortKey="Mugambi, G K" sort="Mugambi, G K" uniqKey="Mugambi G" first="G. K." last="Mugambi">G. K. Mugambi</name><affiliation><nlm:aff id="aff2"><italic>National Museum of Kenya, Botany Department, P.O. Box 40658, 00100, Nairobi, Kenya</italic></nlm:aff></affiliation></author><author><name sortKey="Miller, A N" sort="Miller, A N" uniqKey="Miller A" first="A. N." last="Miller">A. N. Miller</name><affiliation><nlm:aff id="aff3"><italic>Illinois Natural History Survey, University of Illinois Urbana-Champaign, 1816 South Oak Street, Champaign, IL 6182, U.S.A.</italic></nlm:aff></affiliation></author><author><name sortKey="Huhndorf, S M" sort="Huhndorf, S M" uniqKey="Huhndorf S" first="S. M." last="Huhndorf">S. M. Huhndorf</name><affiliation><nlm:aff id="aff4"><italic>The Field Museum, 1400 S. Lake Shore Dr, Chicago, IL 60605, U.S.A.</italic></nlm:aff></affiliation></author><author><name sortKey="Marincowitz, S" sort="Marincowitz, S" uniqKey="Marincowitz S" first="S." last="Marincowitz">S. Marincowitz</name><affiliation><nlm:aff id="aff5"><italic>Forestry and Agricultural Biotechnology Institute, University of Pretoria, Pretoria 0002, South Africa</italic></nlm:aff></affiliation></author><author><name sortKey="Spatafora, J W" sort="Spatafora, J W" uniqKey="Spatafora J" first="J. W." last="Spatafora">J. W. Spatafora</name><affiliation><nlm:aff id="aff6"><italic>Department of Botany and Plant Pathology, Oregon State University, Corvallis, Oregon 93133, U.S.A.</italic></nlm:aff></affiliation></author><author><name sortKey="Schoch, C L" sort="Schoch, C L" uniqKey="Schoch C" first="C. L." last="Schoch">C. L. Schoch</name><affiliation><nlm:aff id="aff7"><italic>National Center for Biotechnology Information (NCBI), National Library of Medicine, National Institutes of Health, GenBank, 45 Center Drive, MSC 6510, Building 45, Room 6an.18, Bethesda, MD, 20892, U.S.A.</italic></nlm:aff></affiliation></author></titleStmt><publicationStmt><idno type="wicri:source">PMC</idno><idno type="pmid">20169023</idno><idno type="pmc">2816966</idno><idno type="url">http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2816966</idno><idno type="RBID">PMC:2816966</idno><idno type="doi">10.3114/sim.2009.64.03</idno><date when="2009">2009</date><idno type="wicri:Area/Pmc/Corpus">000261</idno><idno type="wicri:explorRef" wicri:stream="Pmc" wicri:step="Corpus" wicri:corpus="PMC">000261</idno></publicationStmt><sourceDesc><biblStruct><analytic><title xml:lang="en" level="a" type="main">A molecular phylogenetic reappraisal of the <italic>Hysteriaceae</italic>,
<italic>Mytilinidiaceae</italic> and <italic>Gloniaceae</italic> (<italic>Pleosporomycetidae</italic>,
<italic>Dothideomycetes</italic>) with keys to world species</title><author><name sortKey="Boehm, E W A" sort="Boehm, E W A" uniqKey="Boehm E" first="E. W. A." last="Boehm">E. W. A. Boehm</name><affiliation><nlm:aff id="aff1"><italic>Department of Biological Sciences, Kean University, 1000 Morris Ave., Union, New Jersey 07083, U.S.A.</italic></nlm:aff></affiliation></author><author><name sortKey="Mugambi, G K" sort="Mugambi, G K" uniqKey="Mugambi G" first="G. K." last="Mugambi">G. K. Mugambi</name><affiliation><nlm:aff id="aff2"><italic>National Museum of Kenya, Botany Department, P.O. Box 40658, 00100, Nairobi, Kenya</italic></nlm:aff></affiliation></author><author><name sortKey="Miller, A N" sort="Miller, A N" uniqKey="Miller A" first="A. N." last="Miller">A. N. Miller</name><affiliation><nlm:aff id="aff3"><italic>Illinois Natural History Survey, University of Illinois Urbana-Champaign, 1816 South Oak Street, Champaign, IL 6182, U.S.A.</italic></nlm:aff></affiliation></author><author><name sortKey="Huhndorf, S M" sort="Huhndorf, S M" uniqKey="Huhndorf S" first="S. M." last="Huhndorf">S. M. Huhndorf</name><affiliation><nlm:aff id="aff4"><italic>The Field Museum, 1400 S. Lake Shore Dr, Chicago, IL 60605, U.S.A.</italic></nlm:aff></affiliation></author><author><name sortKey="Marincowitz, S" sort="Marincowitz, S" uniqKey="Marincowitz S" first="S." last="Marincowitz">S. Marincowitz</name><affiliation><nlm:aff id="aff5"><italic>Forestry and Agricultural Biotechnology Institute, University of Pretoria, Pretoria 0002, South Africa</italic></nlm:aff></affiliation></author><author><name sortKey="Spatafora, J W" sort="Spatafora, J W" uniqKey="Spatafora J" first="J. W." last="Spatafora">J. W. Spatafora</name><affiliation><nlm:aff id="aff6"><italic>Department of Botany and Plant Pathology, Oregon State University, Corvallis, Oregon 93133, U.S.A.</italic></nlm:aff></affiliation></author><author><name sortKey="Schoch, C L" sort="Schoch, C L" uniqKey="Schoch C" first="C. L." last="Schoch">C. L. Schoch</name><affiliation><nlm:aff id="aff7"><italic>National Center for Biotechnology Information (NCBI), National Library of Medicine, National Institutes of Health, GenBank, 45 Center Drive, MSC 6510, Building 45, Room 6an.18, Bethesda, MD, 20892, U.S.A.</italic></nlm:aff></affiliation></author></analytic><series><title level="j">Studies in Mycology</title><idno type="ISSN">0166-0616</idno><idno type="eISSN">1872-9797</idno><imprint><date when="2009">2009</date></imprint></series></biblStruct></sourceDesc></fileDesc><profileDesc><textClass></textClass></profileDesc></teiHeader><front><div type="abstract" xml:lang="en"><p>A reappraisal of the phylogenetic integrity of bitunicate ascomycete fungi
belonging to or previously affiliated with the <italic>Hysteriaceae</italic>,
<italic>Mytilinidiaceae</italic>, <italic>Gloniaceae</italic> and <italic>Patellariaceae</italic> is
presented, based on an analysis of 121 isolates and four nuclear genes, the
ribosomal large and small subunits, transcription elongation factor 1 and the
second largest RNA polymerase II subunit. A geographically diverse and high
density taxon sampling strategy was employed, including multiple
isolates/species from the following genera: <italic>Anteaglonium</italic> (6/4),
<italic>Encephalographa</italic> (1/1), <italic>Farlowiella</italic> (3/1),
<italic>Gloniopsis</italic> (8/4), <italic>Glonium</italic> (4/2), <italic>Hysterium</italic> (12/5),
<italic>Hysterobrevium</italic> (14/3), <italic>Hysterographium</italic> (2/1),
<italic>Hysteropatella</italic> (2/2), <italic>Lophium</italic> (4/2), <italic>Mytilinidion</italic>(13/10), <italic>Oedohysterium</italic> (5/3), <italic>Ostreichnion</italic> (2/2),
<italic>Patellaria</italic> (1/1), <italic>Psiloglonium</italic> (11/3), <italic>Quasiconcha</italic>(1/1), <italic>Rhytidhysteron</italic> (8/3), and 24 outgroup taxa. Sequence data
indicate that although the <italic>Hysteriales</italic> are closely related to the
<italic>Pleosporales</italic>, sufficient branch support exists for their separation
into separate orders within the <italic>Pleosporomycetidae</italic>. The
<italic>Mytilinidiales</italic> are more distantly related within the subclass and
show a close association with the <italic>Gloniaceae</italic>. Although there are
examples of concordance between morphological and molecular data, these are
few. Molecular data instead support the premise of a large number of
convergent evolutionary lineages, which do not correspond to previously held
assumptions of synapomorphy relating to spore morphology. Thus, within the
<italic>Hysteriaceae</italic>, the genera <italic>Gloniopsis</italic>, <italic>Glonium</italic>,
<italic>Hysterium</italic> and <italic>Hysterographium</italic> are highly polyphyletic. This
necessitated the transfer of two species of <italic>Hysterium</italic> to
<italic>Oedohysterium</italic> <italic>gen. nov.</italic> (<italic>Od. insidens</italic> <italic>comb.
nov.</italic> and <italic>Od. sinense comb. nov.</italic>), the description of a new
species, <italic>Hysterium barrianum</italic> <italic>sp. nov.</italic>, and the transfer of
two species of <italic>Gloniopsis</italic> to <italic>Hysterobrevium</italic> <italic>gen.
nov.</italic> (<italic>Hb. smilacis</italic> <italic>comb. nov.</italic> and <italic>Hb.
constrictum</italic> <italic>comb. nov.</italic>). While <italic>Hysterographium</italic>, with
the type <italic>Hg. fraxini</italic>, is removed from the <italic>Hysteriaceae</italic>, some
of its species remain within the family, transferred here to
<italic>Oedohysterium</italic> (<italic>Od. pulchrum</italic> <italic>comb. nov.</italic>),
<italic>Hysterobrevium</italic> (<italic>Hb. mori</italic> <italic>comb. nov.</italic>) and
<italic>Gloniopsis</italic> (<italic>Gp. subrugosa</italic> <italic>comb. nov.</italic>); the latter
genus, in addition to the type, <italic>Gp. praelonga</italic>, with two new species,
<italic>Gp. arciformis</italic> <italic>sp. nov.</italic> and <italic>Gp. kenyensis sp. nov</italic>.
The genus <italic>Glonium</italic> is now divided into <italic>Anteaglonium</italic>(<italic>Pleosporales</italic>), <italic>Glonium</italic> (<italic>Gloniaceae</italic>), and
<italic>Psiloglonium</italic> (<italic>Hysteriaceae</italic>). The hysterothecium has evolved
convergently no less than five times within the <italic>Pleosporomycetidae</italic>(e.g., <italic>Anteaglonium</italic>, <italic>Farlowiella</italic>, <italic>Glonium</italic>,
<italic>Hysterographium</italic> and the <italic>Hysteriaceae</italic>). Similarly,
thin-walled mytilinidioid (e.g., <italic>Ostreichnion</italic>) and patellarioid
(e.g., <italic>Rhytidhysteron</italic>) genera, previously in the
<italic>Mytilinidiaceae</italic> and <italic>Patellariaceae</italic>, respectively,
transferred here to the <italic>Hysteriaceae</italic>, have also evolved at least
twice within the subclass. As such, character states traditionally considered
to represent synapomorphies among these fungi, whether they relate to spore
septation or the ascomata, in fact, represent symplesiomorphies, and most
likely have arisen multiple times through convergent evolutionary processes in
response to common selective pressures.</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><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><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><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-id journal-id-type="publisher-id">simycol</journal-id><journal-title-group><journal-title>Studies in Mycology</journal-title></journal-title-group><issn pub-type="ppub">0166-0616</issn><issn pub-type="epub">1872-9797</issn><publisher><publisher-name>CBS Fungal Biodiversity Centre</publisher-name></publisher></journal-meta><article-meta><article-id pub-id-type="pmid">20169023</article-id><article-id pub-id-type="pmc">2816966</article-id><article-id pub-id-type="publisher-id">0049</article-id><article-id pub-id-type="doi">10.3114/sim.2009.64.03</article-id><article-categories><subj-group subj-group-type="heading"><subject>Articles</subject></subj-group></article-categories><title-group><article-title>A molecular phylogenetic reappraisal of the <italic>Hysteriaceae</italic>,
<italic>Mytilinidiaceae</italic> and <italic>Gloniaceae</italic> (<italic>Pleosporomycetidae</italic>,
<italic>Dothideomycetes</italic>) with keys to world species</article-title></title-group><contrib-group><contrib contrib-type="author"><name><surname>Boehm</surname><given-names>E.W.A.</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>Mugambi</surname><given-names>G.K.</given-names></name><xref ref-type="aff" rid="aff2">2</xref></contrib><contrib contrib-type="author"><name><surname>Miller</surname><given-names>A.N.</given-names></name><xref ref-type="aff" rid="aff3">3</xref></contrib><contrib contrib-type="author"><name><surname>Huhndorf</surname><given-names>S.M.</given-names></name><xref ref-type="aff" rid="aff4">4</xref></contrib><contrib contrib-type="author"><name><surname>Marincowitz</surname><given-names>S.</given-names></name><xref ref-type="aff" rid="aff5">5</xref></contrib><contrib contrib-type="author"><name><surname>Spatafora</surname><given-names>J.W.</given-names></name><xref ref-type="aff" rid="aff6">6</xref></contrib><contrib contrib-type="author"><name><surname>Schoch</surname><given-names>C.L.</given-names></name><xref ref-type="aff" rid="aff7">7</xref></contrib></contrib-group><aff id="aff1"><label>1</label><italic>Department of Biological Sciences, Kean University, 1000 Morris Ave., Union, New Jersey 07083, U.S.A.</italic></aff><aff id="aff2"><label>2</label><italic>National Museum of Kenya, Botany Department, P.O. Box 40658, 00100, Nairobi, Kenya</italic></aff><aff id="aff3"><label>3</label><italic>Illinois Natural History Survey, University of Illinois Urbana-Champaign, 1816 South Oak Street, Champaign, IL 6182, U.S.A.</italic></aff><aff id="aff4"><label>4</label><italic>The Field Museum, 1400 S. Lake Shore Dr, Chicago, IL 60605, U.S.A.</italic></aff><aff id="aff5"><label>5</label><italic>Forestry and Agricultural Biotechnology Institute, University of Pretoria, Pretoria 0002, South Africa</italic></aff><aff id="aff6"><label>6</label><italic>Department of Botany and Plant Pathology, Oregon State University, Corvallis, Oregon 93133, U.S.A.</italic></aff><aff id="aff7"><label>7</label><italic>National Center for Biotechnology Information (NCBI), National Library of Medicine, National Institutes of Health, GenBank, 45 Center Drive, MSC 6510, Building 45, Room 6an.18, Bethesda, MD, 20892, U.S.A.</italic></aff><author-notes><corresp id="cor1"><label>*</label><italic>Correspondence</italic>: E.W.A. Boehm,
<email>eboehm@kean.edu</email></corresp></author-notes><pub-date pub-type="ppub"><year>2009</year></pub-date><volume>64</volume><issue-title>A phylogenetic re-evaluation of
<italic>Dothideomycetes</italic></issue-title><fpage>49</fpage><lpage>83-S3</lpage><permissions><copyright-statement>Copyright © Copyright 2009 CBS-KNAW Fungal Biodiversity
Centre</copyright-statement><copyright-year>2009</copyright-year><license><license-p>You are free to share - to copy, distribute and transmit the work, under
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Nothing in this license impairs or restricts the author's moral rights.</license-p></license></permissions><self-uri xlink:title="pdf" xlink:href="49.pdf"></self-uri><abstract><p>A reappraisal of the phylogenetic integrity of bitunicate ascomycete fungi
belonging to or previously affiliated with the <italic>Hysteriaceae</italic>,
<italic>Mytilinidiaceae</italic>, <italic>Gloniaceae</italic> and <italic>Patellariaceae</italic> is
presented, based on an analysis of 121 isolates and four nuclear genes, the
ribosomal large and small subunits, transcription elongation factor 1 and the
second largest RNA polymerase II subunit. A geographically diverse and high
density taxon sampling strategy was employed, including multiple
isolates/species from the following genera: <italic>Anteaglonium</italic> (6/4),
<italic>Encephalographa</italic> (1/1), <italic>Farlowiella</italic> (3/1),
<italic>Gloniopsis</italic> (8/4), <italic>Glonium</italic> (4/2), <italic>Hysterium</italic> (12/5),
<italic>Hysterobrevium</italic> (14/3), <italic>Hysterographium</italic> (2/1),
<italic>Hysteropatella</italic> (2/2), <italic>Lophium</italic> (4/2), <italic>Mytilinidion</italic>(13/10), <italic>Oedohysterium</italic> (5/3), <italic>Ostreichnion</italic> (2/2),
<italic>Patellaria</italic> (1/1), <italic>Psiloglonium</italic> (11/3), <italic>Quasiconcha</italic>(1/1), <italic>Rhytidhysteron</italic> (8/3), and 24 outgroup taxa. Sequence data
indicate that although the <italic>Hysteriales</italic> are closely related to the
<italic>Pleosporales</italic>, sufficient branch support exists for their separation
into separate orders within the <italic>Pleosporomycetidae</italic>. The
<italic>Mytilinidiales</italic> are more distantly related within the subclass and
show a close association with the <italic>Gloniaceae</italic>. Although there are
examples of concordance between morphological and molecular data, these are
few. Molecular data instead support the premise of a large number of
convergent evolutionary lineages, which do not correspond to previously held
assumptions of synapomorphy relating to spore morphology. Thus, within the
<italic>Hysteriaceae</italic>, the genera <italic>Gloniopsis</italic>, <italic>Glonium</italic>,
<italic>Hysterium</italic> and <italic>Hysterographium</italic> are highly polyphyletic. This
necessitated the transfer of two species of <italic>Hysterium</italic> to
<italic>Oedohysterium</italic> <italic>gen. nov.</italic> (<italic>Od. insidens</italic> <italic>comb.
nov.</italic> and <italic>Od. sinense comb. nov.</italic>), the description of a new
species, <italic>Hysterium barrianum</italic> <italic>sp. nov.</italic>, and the transfer of
two species of <italic>Gloniopsis</italic> to <italic>Hysterobrevium</italic> <italic>gen.
nov.</italic> (<italic>Hb. smilacis</italic> <italic>comb. nov.</italic> and <italic>Hb.
constrictum</italic> <italic>comb. nov.</italic>). While <italic>Hysterographium</italic>, with
the type <italic>Hg. fraxini</italic>, is removed from the <italic>Hysteriaceae</italic>, some
of its species remain within the family, transferred here to
<italic>Oedohysterium</italic> (<italic>Od. pulchrum</italic> <italic>comb. nov.</italic>),
<italic>Hysterobrevium</italic> (<italic>Hb. mori</italic> <italic>comb. nov.</italic>) and
<italic>Gloniopsis</italic> (<italic>Gp. subrugosa</italic> <italic>comb. nov.</italic>); the latter
genus, in addition to the type, <italic>Gp. praelonga</italic>, with two new species,
<italic>Gp. arciformis</italic> <italic>sp. nov.</italic> and <italic>Gp. kenyensis sp. nov</italic>.
The genus <italic>Glonium</italic> is now divided into <italic>Anteaglonium</italic>(<italic>Pleosporales</italic>), <italic>Glonium</italic> (<italic>Gloniaceae</italic>), and
<italic>Psiloglonium</italic> (<italic>Hysteriaceae</italic>). The hysterothecium has evolved
convergently no less than five times within the <italic>Pleosporomycetidae</italic>(e.g., <italic>Anteaglonium</italic>, <italic>Farlowiella</italic>, <italic>Glonium</italic>,
<italic>Hysterographium</italic> and the <italic>Hysteriaceae</italic>). Similarly,
thin-walled mytilinidioid (e.g., <italic>Ostreichnion</italic>) and patellarioid
(e.g., <italic>Rhytidhysteron</italic>) genera, previously in the
<italic>Mytilinidiaceae</italic> and <italic>Patellariaceae</italic>, respectively,
transferred here to the <italic>Hysteriaceae</italic>, have also evolved at least
twice within the subclass. As such, character states traditionally considered
to represent synapomorphies among these fungi, whether they relate to spore
septation or the ascomata, in fact, represent symplesiomorphies, and most
likely have arisen multiple times through convergent evolutionary processes in
response to common selective pressures.</p></abstract><kwd-group><kwd>Evolution</kwd><kwd>fungi</kwd><kwd><italic>Hysteriales</italic></kwd><kwd><italic>Mytilinidiales</italic></kwd><kwd><italic>Patellariales</italic></kwd><kwd>phylogeny</kwd><kwd>speciation</kwd><kwd>taxonomy</kwd></kwd-group></article-meta><notes><fn-group><fn><p><bold>Taxonomic novelties:</bold> <bold>New species:</bold> <italic>Gloniopsis
arciformis</italic> E.W.A. Boehm, G.K. Mugambi, S.M. Huhndorf & C.L.
Schoch<italic>, Gp. kenyensis</italic> E.W.A. Boehm, G.K. Mugambi, S.M. Huhndorf &
C.L. Schoch, <italic>Hysterium barrianum</italic> E.W.A. Boehm, A.N. Miller, G.K.
Mugambi, S.M. Huhndorf & C.L. Schoch. <bold>New genera:</bold><italic>Hysterobrevium</italic> E.W.A. Boehm & C.L. Schoch, <italic>Oedohysterium</italic>E.W.A. Boehm & C.L. Schoch. <bold>New combinations:</bold> <italic>Gloniopsis
subrugosa</italic> (Cooke & Ellis) E.W.A. Boehm & C.L. Schoch,
<italic>Hysterobrevium constrictum</italic> (N. Amano) E.W.A. Boehm & C.L. Schoch,
<italic>Hb. mori</italic> (Schwein.) E.W.A. Boehm & C.L. Schoch, <italic>Hb.
smilacis</italic> (Schwein.) E.W.A. Boehm & C.L. Schoch, <italic>Oedohysterium
insidens</italic> (Schwein.) E.W.A. Boehm & C.L. Schoch, <italic>Od. pulchrum</italic>(Checa, Shoemaker & Umaña) E.W.A. Boehm & C.L. Schoch, <italic>Od.
sinense</italic> (Teng) E.W.A. Boehm & C.L. Schoch, <italic>Psiloglonium
araucanum</italic> (Speg.) E.W.A. Boehm, S. Marincowitz & C.L. Schoch, <italic>P.
chambianum</italic> (Guyot) E.W.A. Boehm & C.L. Schoch<italic>, P. colihuae</italic>(Lorenzo & Messuti) E.W.A. Boehm & C.L. Schoch, <italic>P. ephedrae</italic>(Henn.) E.W.A. Boehm & C.L. Schoch<italic>, P. hysterinum</italic> (Rehm) E.W.A.
Boehm & C.L. Schoch<italic>, P. pusillum</italic> (H. Zogg) E.W.A. Boehm &
C.L. Schoch, <italic>P. sasicola</italic> (N. Amano) E.W.A. Boehm & C.L. Schoch,
and <italic>P. uspallatense</italic> (Speg.) E.W.A. Boehm & C.L. Schoch.</p></fn></fn-group></notes></front><body><sec><title>INTRODUCTION</title><p>Class <italic>Dothideomycetes</italic>, subphylum <italic>Pezizomycotina</italic>(<italic>Ascomycota</italic>), is currently classified into two subclasses, based on
centrum type (Schoch <italic>et al.</italic><xref ref-type="bibr" rid="ref100">2006</xref>,
<xref ref-type="bibr" rid="ref101">2009b</xref>,
<xref ref-type="bibr" rid="ref105">Spatafora <italic>et al.</italic>2006</xref>). The <italic>Dothideomycetidae</italic> is characterised by the
absence of sterile centrum elements (<italic>e.g.</italic>, pseudoparaphyses). This
subclass includes the <italic>Dothideales, Capnodiales</italic>, and
<italic>Myriangiales</italic>. The <italic>Microthyriales</italic>, and
<italic>Trypetheliales</italic>, while within the <italic>Dothideomycetes</italic>, lie
outside of the <italic>Dothideomycetidae</italic>(<xref ref-type="bibr" rid="ref99">Schoch <italic>et al.</italic>2009a</xref>). The second subclass recognised within the
<italic>Dothideomycetes</italic> is the <italic>Pleosporomycetidae</italic>, characterised by
a hamathecium of wide to narrow cellular to trabeculate pseudoparaphyses,
which may or may not persist at maturity. This subclass currently comprises
the <italic>Pleosporales, Hysteriales</italic>, and <italic>Mytilinidiales</italic>, and
tentatively the <italic>Jahnulales</italic>. The <italic>Botryosphaeriales</italic>, and
<italic>Patellariales</italic>, possess pseudoparaphyses, and would be expected to
fall into the <italic>Pleosporomycetidae</italic>, however, at present, statistical
support is weak. A greater number of orders, families, and genera still await
placement, and are currently designated as <italic>incertae sedis</italic> within the
<italic>Dothideomycetes</italic> (<xref ref-type="bibr" rid="ref65">Lumbsch &
Huhndorf 2007</xref>).</p><p>Fungi classified in the <italic>Hysteriaceae</italic> (<italic>Hysteriales</italic>),
<italic>Mytilinidiaceae</italic> (<italic>Mytilinidiales</italic>), and <italic>Gloniaceae</italic>(<italic>Pleosporomycetidae fam. incertae sedis</italic>), possess persistent,
carbonaceous ascomata that characteristically dehisce by a longitudinal
suture. Recent molecular data support the inclusion of all three families
within the <italic>Pleosporomycetidae</italic>(<xref ref-type="bibr" rid="ref100">Schoch <italic>et al.</italic> 2006</xref>,
<xref ref-type="bibr" rid="ref18">Boehm <italic>et al.</italic> 2009</xref>,
Mugami & Huhndorf 2009). In the <italic>Hysteriaceae</italic> ascomata are
thick-walled, navicular, characteristically dehiscing by an invaginated slit
or sulcus (<xref ref-type="bibr" rid="ref122">Zogg 1962</xref>). Fungi
in the <italic>Mytilinidiaceae</italic>, on the other hand, possess strongly laterally
compressed, connivent, thin-walled conchate ascomata, reminiscent of miniature
bivalve molluscs. These mytilinidioid ascomata typically dehisce by an
evaginated slit, in some species forming a longitudinal keel or cristate apex
(<xref ref-type="bibr" rid="ref8">Barr 1990a</xref>). Fungi belonging
to the <italic>Gloniaceae</italic>, have dichotomously branched, laterally anastomosed
pseudothecia, that form radiating pseudo-stellate composites and dehisce by an
inconspicuous, longitudinal, but evaginated slit
(<xref ref-type="bibr" rid="ref18">Boehm <italic>et al.</italic>2009</xref>).</p><p>We are broadly interested in the evolution of character states
traditionally used to define higher taxa within each family. Essentially, we
wish to address whether morphological features historically used in the
classification of these fungi are phylogenetically informative in the context
of sequence-based phylogenies. This would have bearing on which morphological
features are phylogenetically significant, and therefore useful for a natural
delineation of higher taxa. Morphological character states traditionally used
to classify these fungi have related primarily to features associated with (1)
the pseudothecium, (2) the peridium, (3) the hamathecium, and (4) differences
in ascospore symmetry (Barr
<xref ref-type="bibr" rid="ref7">1987</xref>,
<xref ref-type="bibr" rid="ref8">1990a</xref>). Character states within
each family relate primarily to ascospore septation and pigmentation
(<xref ref-type="bibr" rid="ref122">Zogg 1962</xref>).</p><p>Due to the seemingly transitional nature of the ascoma, neither fully open
nor closed, hysteriaceous fungi have been placed in the discomycetes and
pyrenomycetes about equally by various mycologists throughout the
19<sup>th</sup> Century (<xref ref-type="bibr" rid="ref14">Bisby
1923</xref>). In his <italic>Systema Mycologicum</italic>, Fries
(<xref ref-type="bibr" rid="ref34">1823</xref>) initially considered
hysteriaceous fungi to belong to the pyrenomycetes and placed them in the
<italic>Phacidiacei</italic>, but later (<xref ref-type="bibr" rid="ref35">Fries
1835</xref>) placed them in his new class <italic>Discomycetes</italic>, stating:
“<italic>Transitum sistunt ad Discomycetes, sed discum verum non
monstrant</italic>.” Chevallier
(<xref ref-type="bibr" rid="ref22">1826</xref>) recognised the unique
nature of the hysterothecium and established the <italic>Hysteriineae</italic>, which
he considered as pyrenomycetes distinct from Fries' <italic>Phacidiei</italic>. Corda
(<xref ref-type="bibr" rid="ref25">1842</xref>), on the other hand,
retained the <italic>Phacidiei</italic> within the <italic>Hysteriaceae</italic>, and divided
the family into a number of subfamilies. De Notaris
(<xref ref-type="bibr" rid="ref28">1847</xref>) considered the
<italic>Hysteriaceae</italic> to belong to the pyrenomycetes and used spore
pigmentation to classify hysteriaceous fungi into the <italic>Phaeosporii</italic> and
the <italic>Hyalosporii</italic>. Saccardo
(<xref ref-type="bibr" rid="ref94">1873</xref>) initially followed
Fries, but later (1874) placed hysteriaceous fungi in the pyrenomycetes, and
carried de Notaris' (<xref ref-type="bibr" rid="ref28">1847</xref>)
spore classification scheme further by dividing the <italic>Hysteriaceae</italic> into
nine sections based on pigmentation and the morphology of spore septation
(<xref ref-type="bibr" rid="ref96">Saccardo 1883</xref>). Ellis &
Everhart (<xref ref-type="bibr" rid="ref32">1892</xref>), in their
<italic>North American Pyrenomycetes</italic>, tentatively included the
<italic>Hysteriaceae</italic>, but stated that they had not at first intended to do so
due to the transitional nature of the hysterothecium. In Rabenhorst's
<italic>Kryptogamen-Flora, Die Pilze</italic>, Rehm
(<xref ref-type="bibr" rid="ref88">1896</xref>) compromised and placed
the <italic>Hysteriales</italic> as an order intermediate between the pyrenomycetes
and the discomycetes.</p><p>Mytilinidioid fungi have also historically been classified within the
family <italic>Hysteriaceae</italic>, due to perceived similarities in ascocarp
morphology, specifically its means of longitudinal dehiscence
(<xref ref-type="bibr" rid="ref34">Fries 1823</xref>,
<xref ref-type="bibr" rid="ref28">De Notaris 1847</xref>, Saccardo
<xref ref-type="bibr" rid="ref95">1875</xref>,
<xref ref-type="bibr" rid="ref96">1883</xref>,
<xref ref-type="bibr" rid="ref32">Ellis & Everhart 1892</xref>,
<xref ref-type="bibr" rid="ref73">Massee 1895</xref>,
<xref ref-type="bibr" rid="ref88">Rehm 1896</xref>,
<xref ref-type="bibr" rid="ref41">von Höhnel 1918</xref>,
<xref ref-type="bibr" rid="ref14">Bisby 1923</xref>). Modern authors
have likewise included mytilinidioid fungi within the <italic>Hysteriaceae</italic>,
placing the family in the <italic>Pseudosphaeriales</italic>(<xref ref-type="bibr" rid="ref83">Nannfeldt 1932</xref>,
<xref ref-type="bibr" rid="ref36">Gäumann 1949</xref>), the
<italic>Dothiorales</italic> (<xref ref-type="bibr" rid="ref82">Müller &
von Arx 1950</xref>, <xref ref-type="bibr" rid="ref2">von Arx &
Müller 1954</xref>), the <italic>Dothideales</italic>(<xref ref-type="bibr" rid="ref3">von Arx & Müller
1975</xref>), and in a separate order <italic>Hysteriales</italic>, closely
related to the <italic>Pleosporales</italic>(<xref ref-type="bibr" rid="ref78">Miller 1949</xref>,
<xref ref-type="bibr" rid="ref69">Luttrell 1955</xref>). The
<italic>Hysteriales</italic> were placed in the subclass <italic>Loculoascomycetes</italic> by
Luttrell (<xref ref-type="bibr" rid="ref69">1955</xref>), due to the
presence of bitunicate asci, corresponding to the <italic>Ascoloculares</italic> first
proposed by Nannfeldt
(<xref ref-type="bibr" rid="ref83">1932</xref>).</p><p>Duby (<xref ref-type="bibr" rid="ref31">1862</xref>) was the first
to propose that hysteriaceous fungi be divided into two sections, the
<italic>Hystériées</italic> and the <italic>Lophiées</italic>, the latter
to accommodate mytilinidioid forms. One hundred years later, Zogg
(<xref ref-type="bibr" rid="ref122">1962</xref>) proposed two families:
the <italic>Hysteriaceae s. str.</italic> to accommodate thick-walled hysteriaceous
forms, and the <italic>Lophiaceae</italic> (<xref ref-type="bibr" rid="ref122">Zogg
1962</xref>, <xref ref-type="bibr" rid="ref3">von Arx & Müller
1975</xref>) to accommodate thin-walled, mytilinidioid fungi. Barr
(<xref ref-type="bibr" rid="ref8">1990a</xref>) made the argument for
retention of the earlier name <italic>Mytilinidiaceae</italic> over the
<italic>Lophiaceae</italic>, despite the proposal to conserve the latter
(<xref ref-type="bibr" rid="ref40">Hawksworth & Eriksson
1988</xref>). Luttrell
(<xref ref-type="bibr" rid="ref68">1953</xref>) studied ascomatal
ontogeny and hamathecial development in <italic>Glonium stellatum</italic>. and
concluded that the <italic>Hysteriaceae</italic> possess the pseudoparaphysate
<italic>Pleospora</italic>-type centrum, in which cellular, septate pseudoparaphyses
grow downwards from the cavity roof, initially anchored at both ends, and
occupy the locule prior to the formation of asci
(<xref ref-type="bibr" rid="ref67">Luttrell 1951</xref>). Luttrell
(<xref ref-type="bibr" rid="ref70">1973</xref>) held a wide concept of
the <italic>Hysteriales</italic>, but did not recognise the family
<italic>Lophiaceae</italic>, instead proposing a subfamily, the <italic>Lophioideae,</italic>within the <italic>Hysteriaceae</italic> to accommodate mytilinidioid forms. Barr
(<xref ref-type="bibr" rid="ref5">1979</xref>) however maintained the
two-family distinction. The <italic>Mytilinidiaceae</italic> was placed in the
<italic>Melanommatales</italic> based on a thin-walled peridium of
scleroparenchymatous cells enclosing a hamathecium of narrow trabeculate
pseudoparaphyses, asci borne in a peripheral layer and with ascospores
typically showing bipolar symmetry (Barr
<xref ref-type="bibr" rid="ref7">1987</xref>,
<xref ref-type="bibr" rid="ref8">1990a</xref>). Later, Barr &
Huhndorf (<xref ref-type="bibr" rid="ref12">2001</xref>) noted that the
family was somewhat atypical of the <italic>Melanommatales</italic>, in that, as a
consequence of reduced locule space attributed to lateral compression, they
possess a basal, rather than peripheral, layer of asci and a reduced
hamathecium at maturity. More recently, the <italic>Melanommatales</italic> have been
included within the <italic>Pleosporales</italic>(<xref ref-type="bibr" rid="ref65">Lumbsch & Huhndorf 2007</xref>).
Barr (<xref ref-type="bibr" rid="ref6">1983</xref>) eventually
abandoned the <italic>Hysteriales</italic> and placed the <italic>Hysteriaceae</italic> within
the <italic>Pleosporales</italic> due to the presence of cellular pseudoparaphyses,
asci borne in a basal rather than peripheral layer and ascospores typically
showing bipolar asymmetry. Eriksson
(<xref ref-type="bibr" rid="ref33">2006</xref>) removed the
<italic>Mytilinidiaceae</italic> from the <italic>Hysteriales</italic> and considered it as
<italic>Dothideomycetes et Chaetothyriomycetes incertae sedis</italic>, leaving the
<italic>Hysteriaceae</italic> as the sole family in the <italic>Hysteriales</italic>.</p><p>More recently, Boehm <italic>et al.</italic>(<xref ref-type="bibr" rid="ref18">2009</xref>) presented the first
combined use of DNA and amino acid sequence data to reconstruct the phylogeny
of hysteriaceous fungi. This study was based on a wide taxon sampling
strategy, and employed four nuclear genes, namely the nuSSU and nuLSU,
Transcription Elongation Factor 1 (<italic>TEF1</italic>) and the second largest RNA
polymerase II subunit (<italic>RPB2</italic>). A number of specific conclusions were
reached: (1) Multigene phylogenies provided strong support for the monophyly
of the <italic>Hysteriaceae</italic> and of the <italic>Mytilinidiaceae</italic>, both within
the <italic>Pleosporomycetidae</italic>. However, sequence data also indicated that
both families were not closely related within the subclass. (2) Although core
groups for many of the genera in the <italic>Hysteriaceae</italic> were defined, the
genera <italic>Hysterium, Gloniopsis</italic>, and <italic>Hysterographium</italic> were
demonstrated to be polyphyletic, with affinities not premised on spore
septation and pigmentation. (3) The genus <italic>Glonium</italic> was also shown to
be polyphyletic, but along two highly divergent lines. <italic>Glonium</italic> lies
outside of the <italic>Hysteriaceae</italic>, and instead finds close affinities with
the family <italic>Mytilinidiaceae</italic>, for which was proposed the
<italic>Gloniaceae</italic> (<xref ref-type="bibr" rid="ref18">Boehm <italic>et al.</italic>2009</xref>), to accommodate the type, <italic>G. stellatum</italic>, and related
forms. (4) The genus <italic>Psiloglonium</italic> was reinstated within the
<italic>Hysteriaceae</italic>, with <italic>P. lineare</italic> as type, to accommodate
didymospored species segregated from <italic>Glonium</italic>. (5) The genera
<italic>Mytilinidion</italic> and <italic>Lophium</italic> formed a strongly supported clade
within the <italic>Pleosporomycetidae</italic>, thus defining the monophyletic
<italic>Mytilinidiaceae</italic>, adjacent to the <italic>Gloniaceae</italic>, for which was
proposed the <italic>Mytilinidiales</italic>(<xref ref-type="bibr" rid="ref18">Boehm <italic>et al.</italic> 2009</xref>).
(6) The genus <italic>Farlowiella</italic>, previously in the <italic>Hysteriaceae</italic>,
was placed as <italic>Pleosporomycetidae gen. incertae sedis.</italic> (7) The genus
<italic>Ostreichnion</italic>, previously in the <italic>Mytilinidiaceae</italic>, was
transferred to the <italic>Hysteriaceae.</italic> (8) The genus
<italic>Rhytidhysteron</italic>, previously in the <italic>Patellariaceae</italic>, was
transferred to the <italic>Hysteriaceae</italic>.</p><p>These taxonomic changes present a number of challenges for understanding
evolution within this group of fungi. The lack of agreement between
morphological character states, previously considered synapomorphic
(<italic>e.g.</italic>, <xref ref-type="bibr" rid="ref122">Zogg 1962</xref>),
and recent molecular data based on the nuSSU, nuLSU, <italic>TEF1</italic> and
<italic>RPB2</italic> (<xref ref-type="bibr" rid="ref18">Boehm <italic>et al.</italic>2009</xref>), had resulted in a highly polyphyletic core set of genera
for the <italic>Hysteriaceae</italic> (<italic>e.g., Hysterium, Hysterographium,
Gloniopsis</italic>, and <italic>Glonium</italic>). This presented us with a complicated
picture of past speciation events for the family, and necessitated the current
reappraisal. Essentially, the challenge was to reconcile discrepancies between
morphological and molecular data, in order to more accurately reflect natural
phylogenetic relationships within the family. As a result, the revised
<italic>Hysteriaceae</italic> bears little resemblance to the original concept of the
family (<xref ref-type="bibr" rid="ref122">Zogg 1962</xref>).</p><p>In an effort to facilitate species identification, a number of dichotomous
keys are presented in the current study. These keys take into consideration
taxonomic changes brought about by DNA and amino acid sequencing studies
(<xref ref-type="bibr" rid="ref100">Schoch <italic>et al.</italic> 2006</xref>,
<xref ref-type="bibr" rid="ref18">Boehm <italic>et al.</italic> 2009</xref>,
<xref ref-type="bibr" rid="ref81">Mugambi & Huhndorf 2009</xref>),
and attempt to provide a morphological basis for the many new relationships
revealed by molecular data. Although the keys are based on those first
presented by Zogg (<xref ref-type="bibr" rid="ref122">1962</xref>),
they considerably expand upon them to include a number of new species and
genera described since the original publication (<italic>e.g.</italic>,
<xref ref-type="bibr" rid="ref26">Darker 1963</xref>,
<xref ref-type="bibr" rid="ref112">Tilak & Kale 1968</xref>, Barr
<xref ref-type="bibr" rid="ref4">1975</xref>,
<xref ref-type="bibr" rid="ref8">1990a</xref>,
<xref ref-type="bibr" rid="ref11">Barr & Blackwell 1980</xref>,
<xref ref-type="bibr" rid="ref1">Amano 1983</xref>,
<xref ref-type="bibr" rid="ref106">Speer 1986</xref>,
<xref ref-type="bibr" rid="ref85">Pande & Rao 1991</xref>,
<xref ref-type="bibr" rid="ref51">van der Linde 1992</xref>,
<xref ref-type="bibr" rid="ref43">Kantvilas & Coppins 1997</xref>,
<xref ref-type="bibr" rid="ref62">Lorenzo & Messuti 1998</xref>,
Messuti & Lorenzo <xref ref-type="bibr" rid="ref75">1997</xref>,
<xref ref-type="bibr" rid="ref76">2003</xref>,
<xref ref-type="bibr" rid="ref77">2007</xref>, Vasilyeva
<xref ref-type="bibr" rid="ref116">2000</xref>,
<xref ref-type="bibr" rid="ref117">2001</xref>,
<xref ref-type="bibr" rid="ref23">Chlebicki & Knudsen 2001</xref>,
<xref ref-type="bibr" rid="ref21">Checa <italic>et al.</italic> 2007</xref>).
In addition to incorporating new species and genera, the revised keys also
take into consideration variation in ascospore measurements as presented by
different authors, and include widened distribution reports as well.
Additional information can be found at
<ext-link ext-link-type="uri" xlink:href="www.eboehm.com/">www.eboehm.com/</ext-link>.</p></sec><sec sec-type="materials|methods"><title>MATERIALS AND METHODS</title><sec><title>Taxon sampling</title><p>Fungal cultures, collection data and DNA GenBank accession numbers are
listed in <xref ref-type="table" rid="tbl1">Table 1</xref> - see online
Supplementary Information. Fungal cultures initiated for this study were based
on the isolation of individual ascospores, employing a method whereby
individual ascomata were affixed to Petri plate lids suspended over
potato-dextrose agar. Every 12 h the lids were rotated 45 degrees, such that
after 96 h, confirmation of spore deposits could be made under a
stereomicroscope using transmitted light. Discharged spores were observed
microscopically to confirm identity, transferring a single ascospore to
initiate an axenic culture (<italic>e.g.</italic>, EB cultures and deposits with the
CBS; Centraalbureau voor Schimmelcultures). In some cases, spore discharge was
not obtained, necessitating DNA extraction from individual fruitbodies
(<italic>e.g.</italic>, all GKM, SMH, ANM and some EB accessions). Lastly, a number of
original cultures, from the CBS were employed in this study, the provenance of
which could not be ascertained beforehand. Confirmation of taxonomic identity
was based on whether different isolates of the same species co-segregated in
the final tree.</p><p><table-wrap position="float" id="tbl1"><label>Table 1.</label><caption><p>Taxon sampling, provenance and GenBank accession numbers.</p></caption><table frame="hsides" rules="groups"><thead><tr><th align="left" valign="top"><bold>Species</bold></th><th align="left" valign="top"><bold>Accession</bold></th><th align="left" valign="top"><bold>Provenance</bold></th><th colspan="4" align="left" valign="top"><bold>Genbank No.</bold></th></tr><tr><th valign="top" align="left"></th><th valign="top" align="left"></th><th valign="top" align="left"></th><th valign="top" align="left"><bold>nuSSU</bold></th><th valign="top" align="left"><bold>nuLSU</bold></th><th valign="top" align="left"><bold><italic>TEF1</italic></bold></th><th valign="top" align="left"><bold><italic>RPB2</italic></bold></th></tr></thead><tbody><tr><td align="left" valign="top"><italic>Acrogenospora sphaerocephala</italic></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=164.76&link_type=cbs">CBS 164.76</ext-link></bold></td><td align="left" valign="top"> W. Gams, Grande Tinémont, BELGIUM
</td><td align="left" valign="top"><bold>GU296129</bold></td><td align="left" valign="top"><bold>GU301791</bold></td><td align="left" valign="top"><bold>GU349059</bold></td><td align="left" valign="top"><bold>GU371748</bold></td></tr><tr><td align="left" valign="top"><italic>Aliquandostipite khaoyaiensis</italic></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=118232&link_type=cbs">CBS 118232</ext-link></bold></td><td align="left" valign="top"> P. Inderbitzin (AFTOL 1364), Khao Yai NP, THAILAND
</td><td align="left" valign="top"> AF201453
</td><td align="left" valign="top"><bold>GU301796</bold></td><td align="left" valign="top"><bold>GU349048</bold></td><td align="left" valign="top"> FJ238360
</td></tr><tr><td align="left" valign="top"><italic>Anteaglonium abbreviatum</italic></td><td align="left" valign="top"><bold>ANM 925.1</bold></td><td align="left" valign="top"> A.N. Miller (ILLS), Smoky Mts. TN, U.S.A.
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> GQ221877
</td><td align="left" valign="top"> GQ221924
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"><italic>A. globosum</italic></td><td align="left" valign="top"><bold>SMH 5283</bold></td><td align="left" valign="top"> S.M. Huhndorf (F), Indiana Dunes, IN, U.S.A.
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> GQ221911
</td><td align="left" valign="top"> GQ221919
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"></td><td align="left" valign="top"><bold>ANM 925.2</bold></td><td align="left" valign="top"> A.N. Miller (ILLS), Smoky Mts. TN, U.S.A.
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> GQ221879
</td><td align="left" valign="top"> GQ221925
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"><italic>A. latirostrum</italic></td><td align="left" valign="top"><bold>GKM L100N.2</bold></td><td align="left" valign="top"> G.K. Mugambi (EA), Taita Hills, KENYA
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> GQ221876
</td><td align="left" valign="top"> GQ221938
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"></td><td align="left" valign="top"><bold>GKM 1119</bold></td><td align="left" valign="top"> G.K. Mugambi (EA), Taita Hills, KENYA
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> GQ221874
</td><td align="left" valign="top"> GQ221937
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"><italic>A. parvulum</italic></td><td align="left" valign="top"><bold>SMH 5210</bold></td><td align="left" valign="top"> S.M. Huhndorf (F), NEW ZEALAND
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> GQ221907
</td><td align="left" valign="top"> GQ221917
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"><italic>Arthonia caesia</italic></td><td align="left" valign="top"><bold>AFTOL 775</bold></td><td align="left" valign="top"> A. Amtoft, NC, U.S.A.
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> FJ469668
</td><td align="left" valign="top"> FJ469669
</td><td align="left" valign="top"> FJ469670
</td></tr><tr><td align="left" valign="top"><italic>Botryosphaeria dothidea</italic></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=115476&link_type=cbs">CBS 115476</ext-link></bold></td><td align="left" valign="top"> B. Slippers (AFTOL 946), Crocifisso, SWITZERLAND
</td><td align="left" valign="top"> DQ677998
</td><td align="left" valign="top"> DQ678051
</td><td align="left" valign="top"> DQ767637
</td><td align="left" valign="top"> DQ677944
</td></tr><tr><td align="left" valign="top"><italic>Byssothecium circinans</italic></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=675.92&link_type=cbs">CBS 675.92</ext-link></bold></td><td align="left" valign="top"> G. Semeniuk (AFTOL 1735), SD, U.S.A.
</td><td align="left" valign="top"> AY016339
</td><td align="left" valign="top"> AY016357
</td><td align="left" valign="top"> GU349061
</td><td align="left" valign="top"> DQ767646
</td></tr><tr><td align="left" valign="top"><italic>Cenococcum geophilum</italic></td><td align="left" valign="top"><bold>HUNT A1</bold></td><td align="left" valign="top"> K.F. LoBuglio, GenBank
</td><td align="left" valign="top"> L76616
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"><italic>C. geophilum</italic></td><td align="left" valign="top"><bold>CGMONT</bold></td><td align="left" valign="top"> K.F. LoBuglio, GenBank
</td><td align="left" valign="top"> L76617
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"></td><td align="left" valign="top"><bold>010</bold></td><td align="left" valign="top"> K.F. LoBuglio, GenBank
</td><td align="left" valign="top"> L76618
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"><italic>Cochliobolus heterostrophus</italic></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=134.39&link_type=cbs">CBS 134.39</ext-link></bold></td><td align="left" valign="top"> K. Böning (AFTOL 54)
</td><td align="left" valign="top"> AY544727
</td><td align="left" valign="top"> AY544645
</td><td align="left" valign="top"> DQ497603
</td><td align="left" valign="top"> DQ247790
</td></tr><tr><td align="left" valign="top"><italic>Delitschia winteri</italic></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=225.62&link_type=cbs">CBS 225.62</ext-link></bold></td><td align="left" valign="top"> J.L. Bezerra (AFTOL 1599), Baarn, NETHERLANDS
</td><td align="left" valign="top"> DQ678026
</td><td align="left" valign="top"> DQ678077
</td><td align="left" valign="top"> DQ677922
</td><td align="left" valign="top"> DQ677975
</td></tr><tr><td align="left" valign="top"><italic>Dothidea insculpta</italic></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=189.58&link_type=cbs">CBS 189.58</ext-link></bold></td><td align="left" valign="top"> E. Müller (AFTOL921), Maupas, FRANCE
</td><td align="left" valign="top"> DQ247810
</td><td align="left" valign="top"> DQ247802
</td><td align="left" valign="top"> DQ471081
</td><td align="left" valign="top"> AF107800
</td></tr><tr><td align="left" valign="top"><italic>D. sambuci</italic></td><td align="left" valign="top"><bold>DAOM 231303</bold></td><td align="left" valign="top"> S. Hambleton & B. Shoemaker (AFTOL 274)
</td><td align="left" valign="top"> AY544722
</td><td align="left" valign="top"> AY544681
</td><td align="left" valign="top"> DQ497606
</td><td align="left" valign="top"> DQ522854
</td></tr><tr><td align="left" valign="top"><italic>Elsinoë veneta</italic></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=150.27&link_type=cbs">CBS 150.27</ext-link></bold></td><td align="left" valign="top"> E.M. Wakefield (AFTOL 1853)
</td><td align="left" valign="top"> DQ767651
</td><td align="left" valign="top"> DQ767658
</td><td align="left" valign="top"> DQ767641
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"><italic>Encephalographa elisae</italic></td><td align="left" valign="top"><bold>EB 0347</bold></td><td align="left" valign="top"> M. Tretiach, (BPI 879773), Prov. Trieste, Karst, ITALY
</td><td align="left" valign="top"><bold>GU397358</bold></td><td align="left" valign="top"><bold>GU397343</bold></td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"><italic>Farlowiella carmichaeliana</italic></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=206.36&link_type=cbs">CBS 206.36</ext-link></bold></td><td align="left" valign="top"> E.W. Mason (AFTOL1787). EUROPE
</td><td align="left" valign="top"> AY541482
</td><td align="left" valign="top"> AY541492
</td><td align="left" valign="top"> DQ677931
</td><td align="left" valign="top"> DQ677989
</td></tr><tr><td align="left" valign="top"><italic>F. carmichaeliana</italic></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=179.73&link_type=cbs">CBS 179.73</ext-link></bold></td><td align="left" valign="top"> W. Gams, Teutoburger Wald, Neuenheerse, GERMANY
</td><td align="left" valign="top"> GU296148
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"><italic>Gloniopsis arciformis</italic></td><td align="left" valign="top"><bold>GKM L166A</bold></td><td align="left" valign="top"> G.K. Mugambi (BPI 879774 = Holotype), Malindi, KENYA
</td><td align="left" valign="top"><bold>GU323180</bold></td><td align="left" valign="top"><bold>GU323211</bold></td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"><italic>Gp. kenyensis</italic></td><td align="left" valign="top"><bold>GKM 1010</bold></td><td align="left" valign="top"> G.K. Mugambi (BPI 879775 = Holotype), EA, Malindi, KENYA
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> GQ221891
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"><italic>Gp. praelonga</italic></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=112415&link_type=cbs">CBS 112415</ext-link></bold></td><td align="left" valign="top"> S. Marincowitz (PREM), Kogelberg NR, SOUTH AFRICA
</td><td align="left" valign="top"> FJ161134
</td><td align="left" valign="top"> FJ161173
</td><td align="left" valign="top"> FJ161090
</td><td align="left" valign="top"> FJ161113
</td></tr><tr><td align="left" valign="top"></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123337&link_type=cbs">CBS 123337</ext-link></bold></td><td align="left" valign="top"> E.W.A. Boehm (BPI 878725), NJ, U.S.A.
</td><td align="left" valign="top"> FJ161154
</td><td align="left" valign="top"> FJ161195
</td><td align="left" valign="top"> FJ161103
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"></td><td align="left" valign="top"><bold>CMW 19983</bold></td><td align="left" valign="top"> S. Marincowitz (PREM 57539), Jonkershoek, SOUTH AFRICA
</td><td align="left" valign="top"> FJ161152
</td><td align="left" valign="top"> FJ161193
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"><italic>Gp. subrugosa</italic></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123346&link_type=cbs">CBS 123346</ext-link></bold></td><td align="left" valign="top"> S. Marincowitz (BPI 878735), Gauteng, SOUTH AFRICA
</td><td align="left" valign="top"> FJ161170
</td><td align="left" valign="top"> FJ161210
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> FJ161131
</td></tr><tr><td align="left" valign="top"></td><td align="left" valign="top"><bold>GKM 1214</bold></td><td align="left" valign="top"> G.K. Mugambi (BPI 879776, EA), Mt. Kenya, KENYA
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> GQ221895
</td><td align="left" valign="top"><bold>GU397336</bold></td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"></td><td align="left" valign="top"><bold>SMH 557</bold></td><td align="left" valign="top"> S.M. Huhndorf (BPI 879777, F), Sancti Spiritus, CUBA
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> GQ221896
</td><td align="left" valign="top"><bold>GU397337</bold></td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"><italic>Glonium circumserpens</italic></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123342&link_type=cbs">CBS 123342</ext-link></bold></td><td align="left" valign="top"> G. Kantvilas (BPI 878738), Warra SST, TASMANIA
</td><td align="left" valign="top"> FJ161168
</td><td align="left" valign="top"> FJ161208
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"><italic>G. circumserpens</italic></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123343&link_type=cbs">CBS 123343</ext-link></bold></td><td align="left" valign="top"> G. Kantvilas (BPI 878739), Warra SST, TASMANIA
</td><td align="left" valign="top"> FJ161160
</td><td align="left" valign="top"> FJ161200
</td><td align="left" valign="top"> FJ161108
</td><td align="left" valign="top"> FJ161126
</td></tr><tr><td align="left" valign="top"><italic>G. stellatum</italic></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=207.34&link_type=cbs">CBS 207.34</ext-link></bold></td><td align="left" valign="top"> M.L. Lohman (No. 265), MI, U.S.A.
</td><td align="left" valign="top"> FJ161140
</td><td align="left" valign="top"> FJ161179
</td><td align="left" valign="top"> FJ161095
</td><td align="left" valign="top"></td></tr><tr><td align="left" valign="top"></td><td align="left" valign="top"><bold>ANM 32</bold></td><td align="left" valign="top"> A.N. Miller (ILLS), Smoky Mts., TN, U.S.A.
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> GQ221887
</td><td align="left" valign="top"></td><td align="left" valign="top"></td></tr><tr><td align="left" valign="top"><italic>Guignardia gaultheriae</italic></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=447.70&link_type=cbs">CBS 447.70</ext-link></bold></td><td align="left" valign="top"> H.A. van der Aa (AFTOL 1784), Seattle, WA, U.S.A.
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> DQ678089
</td><td align="left" valign="top"></td><td align="left" valign="top"> DQ677987
</td></tr><tr><td align="left" valign="top"><italic>Herpotrichia diffusa</italic></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=250.62&link_type=cbs">CBS 250.62</ext-link></bold></td><td align="left" valign="top"> M.C. Pande (AFTOL 1588), Uttar Pradesh, INDIA
</td><td align="left" valign="top"> DQ678019
</td><td align="left" valign="top"> DQ678071
</td><td align="left" valign="top"> DQ677915
</td><td align="left" valign="top"> DQ677968
</td></tr><tr><td align="left" valign="top"><italic>Hysterium angustatum</italic></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=236.34&link_type=cbs">CBS 236.34</ext-link></bold></td><td align="left" valign="top"> M.L. Lohman (No. 309), WI, U.S.A.
</td><td align="left" valign="top"><bold>GU397359</bold></td><td align="left" valign="top"> FJ161180
</td><td align="left" valign="top"> FJ161096
</td><td align="left" valign="top"> FJ161117
</td></tr><tr><td align="left" valign="top"></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123334&link_type=cbs">CBS 123334</ext-link></bold></td><td align="left" valign="top"> E.W.A. Boehm (BPI 878724), Sussex Co., NJ, U.S.A.
</td><td align="left" valign="top"> FJ161167
</td><td align="left" valign="top"> FJ161207
</td><td align="left" valign="top"> FJ161111
</td><td align="left" valign="top"> FJ161129
</td></tr><tr><td align="left" valign="top"></td><td align="left" valign="top"><bold>CMW 20409</bold></td><td align="left" valign="top"> S. Marincowitz (PREM 57585), Kleinmond, SOUTH AFRICA
</td><td align="left" valign="top"> FJ161153
</td><td align="left" valign="top"> FJ161194
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"></td><td align="left" valign="top"><bold>SMH 5211.0</bold></td><td align="left" valign="top"> S.M. Huhndorf (F), NEW ZEALAND
</td><td align="left" valign="top"><bold>GU397360</bold></td><td align="left" valign="top"> GQ221905
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> GQ221923
</td></tr><tr><td align="left" valign="top"></td><td align="left" valign="top"><bold>GKM 243A</bold></td><td align="left" valign="top"> G.K. Mugambi (EA), Malindi, KENYA
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> GQ221899
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"></td><td align="left" valign="top"><bold>SMH 5216</bold></td><td align="left" valign="top"> S.M. Huhndorf (F), NEW ZEALAND
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> GQ221933
</td></tr><tr><td align="left" valign="top"></td><td align="left" valign="top"><bold>ANM 85</bold></td><td align="left" valign="top"> A.N. Miller (ILLS), Smoky Mts., TN, U.S.A.
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> GQ221898
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"><italic>H. barrianum</italic></td><td align="left" valign="top"><bold>ANM 1495</bold></td><td align="left" valign="top"> A.N. Miller (ILLS59908 = Holotype, BPI879783 = Paratype), TN, U.S.A.
</td><td align="left" valign="top"><bold>GU323182</bold></td><td align="left" valign="top"> GQ221885
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"></td><td align="left" valign="top"><bold>ANM 1442</bold></td><td align="left" valign="top"> A.N. Miller (ILLS 59907, BPI 879784), Smoky Mts., TN, U.S.A.
</td><td align="left" valign="top"><bold>GU323181</bold></td><td align="left" valign="top"> GQ221884
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"><italic>H. hyalinum</italic></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=237.34&link_type=cbs">CBS 237.34</ext-link></bold></td><td align="left" valign="top"> M.L. Lohman (No. 425), MA, U.S.A.
</td><td align="left" valign="top"> FJ161141
</td><td align="left" valign="top"> FJ161181
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"><italic>H. pulicare</italic></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123377&link_type=cbs">CBS 123377</ext-link></bold></td><td align="left" valign="top"> E.W.A. Boehm (BPI 878723), NY, U.S.A.
</td><td align="left" valign="top"> FJ161161
</td><td align="left" valign="top"> FJ161201
</td><td align="left" valign="top"> FJ161109
</td><td align="left" valign="top"> FJ161127
</td></tr><tr><td align="left" valign="top"><italic>H. vermiforme</italic></td><td align="left" valign="top"><bold>GKM 1234</bold></td><td align="left" valign="top"> G.K. Mugambi (BPI 879785, EA), Mt. Kenya, KENYA
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> GQ221897
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"><italic>Hysterobrevium constrictum</italic></td><td align="left" valign="top"><bold>SMH 5211.1</bold></td><td align="left" valign="top"> S.M. Huhndorf (F), NEW ZEALAND
</td><td align="left" valign="top"><bold>GU397361</bold></td><td align="left" valign="top"> GQ221905
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"><italic>Hb. mori</italic></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=245.34&link_type=cbs">CBS 245.34</ext-link></bold></td><td align="left" valign="top"> M.L. Lohman (No. 6), MI, U.S.A.
</td><td align="left" valign="top"> FJ161143
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> FJ161098
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123563&link_type=cbs">CBS 123563</ext-link></bold></td><td align="left" valign="top"> E.W.A. Boehm (BPI 878731), NY, U.S.A.
</td><td align="left" valign="top"> FJ161155
</td><td align="left" valign="top"> FJ161196
</td><td align="left" valign="top"> FJ161104
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123564&link_type=cbs">CBS 123564</ext-link></bold></td><td align="left" valign="top"> E.W.A. Boehm (BPI 878732), NJ, U.S.A.
</td><td align="left" valign="top"> FJ161158
</td><td align="left" valign="top"> FJ161198
</td><td align="left" valign="top"> FJ161106
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123336&link_type=cbs">CBS 123336</ext-link></bold></td><td align="left" valign="top"> E.W.A. Boehm (BPI 878733), NJ, U.S.A.
</td><td align="left" valign="top"> FJ161164
</td><td align="left" valign="top"> FJ161204
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123335&link_type=cbs">CBS 123335</ext-link></bold></td><td align="left" valign="top"> E.W.A. Boehm (BPI 878734), NY, U.S.A.
</td><td align="left" valign="top"> FJ161162
</td><td align="left" valign="top"> FJ161202
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"></td><td align="left" valign="top"><bold>SMH 5273</bold></td><td align="left" valign="top"> S.M. Huhndorf (BPI 879787, F), IN, U.S.A.
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> GQ221910
</td><td align="left" valign="top"> GQ221936
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"></td><td align="left" valign="top"><bold>GKM 1013</bold></td><td align="left" valign="top"> G.K. Mugambi (BPI 879788, EA), Malindi, KENYA
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"><bold>GU397344</bold></td><td align="left" valign="top"><bold>GU397338</bold></td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"></td><td align="left" valign="top"><bold>SMH 5286</bold></td><td align="left" valign="top"> G.K. Mugambi (BPI 879789, EA), MI, U.S.A.
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"><bold>GU397345</bold></td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"><italic>Hb. smilacis</italic></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=114601&link_type=cbs">CBS 114601</ext-link></bold></td><td align="left" valign="top"> O. Constantinescu, as <italic>Gp. curvata</italic> (Fr.) Sacc., SWEDEN
</td><td align="left" valign="top"> FJ161135
</td><td align="left" valign="top"> FJ161174
</td><td align="left" valign="top"> FJ161091
</td><td align="left" valign="top"> FJ161114
</td></tr><tr><td align="left" valign="top"></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=200.34&link_type=cbs">CBS 200.34</ext-link></bold></td><td align="left" valign="top"> M.L. Lohman (No. 29), as Gp. gerardiana Sacc., MI, U.S.A.
</td><td align="left" valign="top"> FJ161138
</td><td align="left" valign="top"> FJ161177
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"></td><td align="left" valign="top"><bold>CMW 18053</bold></td><td align="left" valign="top"> S. Marincowitz (PREM 57546), Kirstenbosch, SOUTH AFRICA
</td><td align="left" valign="top"> FJ161150
</td><td align="left" valign="top"> FJ161191
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"></td><td align="left" valign="top"><bold>SMH 5280</bold></td><td align="left" valign="top"> G.K. Mugambi (EA), IN, U.S.A.
</td><td align="left" valign="top"><bold>GU323183</bold></td><td align="left" valign="top"> GQ221912
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"><bold>GU371784</bold></td></tr><tr><td align="left" valign="top"></td><td align="left" valign="top"><bold>GKM 426N</bold></td><td align="left" valign="top"> G.K. Mugambi (EA), Taita Hills, KENYA
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> GQ221901
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"><italic>Hysterographium fraxini</italic></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=109.43&link_type=cbs">CBS 109.43</ext-link></bold></td><td align="left" valign="top"> H. Zogg, SWITZERLAND
</td><td align="left" valign="top"> FJ161132
</td><td align="left" valign="top"> FJ161171
</td><td align="left" valign="top"> FJ161088
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"><italic>Hg. fraxini</italic></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=242.34&link_type=cbs">CBS 242.34</ext-link></bold></td><td align="left" valign="top"> M.L. Lohman (No. 300), Manitoba, CANADA
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> FJ161189
</td><td align="left" valign="top"></td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"><italic>Hysteropatella clavispora</italic></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=247.34&link_type=cbs">CBS 247.34</ext-link></bold></td><td align="left" valign="top"> M.L. Lohman (No. 143), IN, U.S.A.
</td><td align="left" valign="top"> DQ678006
</td><td align="left" valign="top"> AY541493
</td><td align="left" valign="top"> DQ677901
</td><td align="left" valign="top"> DQ677955
</td></tr><tr><td align="left" valign="top"><italic>Hp. elliptica</italic></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=935.97&link_type=cbs">CBS 935.97</ext-link></bold></td><td align="left" valign="top"> G. Marson (AFTOL 1790), Fentange, LUXEMBOURG
</td><td align="left" valign="top"> EF495114
</td><td align="left" valign="top"> DQ767657
</td><td align="left" valign="top"> DQ767640
</td><td align="left" valign="top"> DQ767647
</td></tr><tr><td align="left" valign="top"><italic>Jahnula aquatica</italic></td><td align="left" valign="top"><bold>R68-1</bold></td><td align="left" valign="top"> Campbell <italic>et al.</italic> 2007
</td><td align="left" valign="top"> EF175633
</td><td align="left" valign="top"> DF175655
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"><italic>Leptosphaeria maculans</italic></td><td align="left" valign="top"><bold>DAOM 229267</bold></td><td align="left" valign="top"> S. Hambleton & B. Shoemaker (AFTOL 277), CANADA
</td><td align="left" valign="top"> DQ470993
</td><td align="left" valign="top"> DQ470946
</td><td align="left" valign="top"> DQ471062
</td><td align="left" valign="top"> DQ470894
</td></tr><tr><td align="left" valign="top"><italic>Lophium elegans</italic></td><td align="left" valign="top"><bold>EB 0366</bold></td><td align="left" valign="top"> A. Gardiennet (BPI 879792), Til-Chatel, FRANCE
</td><td align="left" valign="top"><bold>GU323184</bold></td><td align="left" valign="top"><bold>GU323210</bold></td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"><italic>L. mytilinum</italic></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=269.34&link_type=cbs">CBS 269.34</ext-link></bold></td><td align="left" valign="top"> M.L. Lohman (AFTOL 1609), MI, U.S.A.
</td><td align="left" valign="top"> DQ678030
</td><td align="left" valign="top"> DQ678081
</td><td align="left" valign="top"> DQ677926
</td><td align="left" valign="top"> DQ677979
</td></tr><tr><td align="left" valign="top"></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=114111&link_type=cbs">CBS 114111</ext-link></bold></td><td align="left" valign="top"> K. & L. Holm & O Constantinescu, Uppland, SWEDEN
</td><td align="left" valign="top"> EF596819
</td><td align="left" valign="top"> EF596819
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123344&link_type=cbs">CBS 123344</ext-link></bold></td><td align="left" valign="top"> E.W.A. Boehm (BPI 878736), NY, U.S.A.
</td><td align="left" valign="top"> FJ161163
</td><td align="left" valign="top"> FJ161203
</td><td align="left" valign="top"> FJ161110
</td><td align="left" valign="top"> FJ161128
</td></tr><tr><td align="left" valign="top"><italic>Mycosphaerella punctiformis</italic></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=113265&link_type=cbs">CBS 113265</ext-link></bold></td><td align="left" valign="top"> G. Verkley (AFTOL 942), Utrecht, NETHERLANDS
</td><td align="left" valign="top"> DQ471017
</td><td align="left" valign="top"> DQ470968
</td><td align="left" valign="top"> DQ471092
</td><td align="left" valign="top"> DQ470920
</td></tr><tr><td align="left" valign="top"><italic>Myriangium duriaei</italic></td><td align="left" valign="top"><bold>AFTOL 1304</bold></td><td align="left" valign="top"> L. Grodsinsky (<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=260.36&link_type=cbs">CBS
260.36</ext-link>), Delta del Parana, ARGENTINA
</td><td align="left" valign="top"> AY016347
</td><td align="left" valign="top"> DQ678059
</td><td align="left" valign="top"> DQ677900
</td><td align="left" valign="top"> DQ677954
</td></tr><tr><td align="left" valign="top"><italic>Mytilinidion acicola</italic></td><td align="left" valign="top"><bold>EB 0379</bold></td><td align="left" valign="top"> A. Gardiennet (BPI 879793), Veronnes, FRANCE
</td><td align="left" valign="top"><bold>GU397362</bold></td><td align="left" valign="top"><bold>GU397346</bold></td><td align="left" valign="top"> —
</td><td align="left" valign="top"><bold>GU397355</bold></td></tr><tr><td align="left" valign="top"><italic>M. acicola</italic></td><td align="left" valign="top"><bold>EB 0349</bold></td><td align="left" valign="top"> A. Gardiennet (BPI 879794), Fixey, Combe Laveau, FRANCE
</td><td align="left" valign="top"><bold>GU323185</bold></td><td align="left" valign="top"><bold>GU323209</bold></td><td align="left" valign="top"> —
</td><td align="left" valign="top"><bold>GU371757</bold></td></tr><tr><td align="left" valign="top"><italic>M. andinense</italic></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123562&link_type=cbs">CBS 123562</ext-link></bold></td><td align="left" valign="top"> M.I. Messuti (BPI 878737), Barrio Don Orione, ARGENTINA.
</td><td align="left" valign="top"> FJ161159
</td><td align="left" valign="top"> FJ161199
</td><td align="left" valign="top"> FJ161107
</td><td align="left" valign="top"> FJ161125
</td></tr><tr><td align="left" valign="top"><italic>M. australe</italic></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=301.34&link_type=cbs">CBS 301.34</ext-link></bold></td><td align="left" valign="top"> A.H. Smith & M.L. Lohman, (type culture), LA, U.S.A.
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> FJ161183
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"><italic>M. californicum</italic></td><td align="left" valign="top"><bold>EB 0385</bold></td><td align="left" valign="top"> A. Gardiennet (BPI 879795), Bois de la Chamage, FRANCE
</td><td align="left" valign="top"><bold>GU323186</bold></td><td align="left" valign="top"><bold>GU323208</bold></td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"><italic>M. mytilinellum</italic></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=303.34&link_type=cbs">CBS 303.34</ext-link></bold></td><td align="left" valign="top"> M.L. Lohman (No. 281), as M. laeviusculum, MI, U.S.A.
</td><td align="left" valign="top"> FJ161144
</td><td align="left" valign="top"> FJ161184
</td><td align="left" valign="top"> FJ161100
</td><td align="left" valign="top"> FJ161119
</td></tr><tr><td align="left" valign="top"></td><td align="left" valign="top"><bold>EB 0386</bold></td><td align="left" valign="top"> A. Gardiennet (BPI 879796), Boissenois, FRANCE
</td><td align="left" valign="top"><bold>GU397363</bold></td><td align="left" valign="top"><bold>GU397347</bold></td><td align="left" valign="top"> —
</td><td align="left" valign="top"><bold>GU397356</bold></td></tr><tr><td align="left" valign="top"><italic>M. resinicola</italic></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=304.34&link_type=cbs">CBS 304.34</ext-link></bold></td><td align="left" valign="top"> M.L. Lohman, No. 260, MI, U.S.A.
</td><td align="left" valign="top"> FJ161145
</td><td align="left" valign="top"> FJ161185
</td><td align="left" valign="top"> FJ161101
</td><td align="left" valign="top"> FJ161120
</td></tr><tr><td align="left" valign="top"><italic>M. rhenanum</italic></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=135.45&link_type=cbs">CBS 135.45</ext-link></bold></td><td align="left" valign="top"> NCTC 6434 (1945), as <italic>M. karstenii</italic></td><td align="left" valign="top"> FJ161136
</td><td align="left" valign="top"> FJ161175
</td><td align="left" valign="top"> FJ161092
</td><td align="left" valign="top"> FJ161115
</td></tr><tr><td align="left" valign="top"></td><td align="left" valign="top"><bold>EB 0341</bold></td><td align="left" valign="top"> A. Brissard, Guesnes, FRANCE
</td><td align="left" valign="top"><bold>GU323187</bold></td><td align="left" valign="top"><bold>GU323207</bold></td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"><italic>M. scolecosporum</italic></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=305.34&link_type=cbs">CBS 305.34</ext-link></bold></td><td align="left" valign="top"> A.H. Smith & M.L. Lohman, WI, U.S.A.
</td><td align="left" valign="top"> FJ161146
</td><td align="left" valign="top"> FJ161186
</td><td align="left" valign="top"> FJ161102
</td><td align="left" valign="top"> FJ161121
</td></tr><tr><td align="left" valign="top"><italic>M. thujarum</italic></td><td align="left" valign="top"><bold>EB 0268</bold></td><td align="left" valign="top"> E.W.A. Boehm (BPI 879797), NY, U.S.A.
</td><td align="left" valign="top"><bold>GU323188</bold></td><td align="left" valign="top"><bold>GU323206</bold></td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"><italic>M. tortile</italic></td><td align="left" valign="top"><bold>EB 0377</bold></td><td align="left" valign="top"> A. Gardiennet (BPI 879798), Veronnes, FRANCE
</td><td align="left" valign="top"><bold>GU323189</bold></td><td align="left" valign="top"><bold>GU323205</bold></td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"><italic>Oedohysterium insidens</italic></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=238.34&link_type=cbs">CBS 238.34</ext-link></bold></td><td align="left" valign="top"> M.L. Lohman (No. 308) MI, U.S.A.
</td><td align="left" valign="top"> FJ161142
</td><td align="left" valign="top"> FJ161182
</td><td align="left" valign="top"> FJ161097
</td><td align="left" valign="top"> FJ161118
</td></tr><tr><td align="left" valign="top"><italic>Od. insidens</italic></td><td align="left" valign="top"><bold>ANM 1443</bold></td><td align="left" valign="top"> A.N. Miller (BPI 879799, ILLS), Smoky Mts., TN, U.S.A.
</td><td align="left" valign="top"><bold>GU323190</bold></td><td align="left" valign="top"> GQ221882
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"><bold>GU371785</bold></td></tr><tr><td align="left" valign="top"><italic>Od. pulchrum</italic></td><td align="left" valign="top"><bold>DQ 402184</bold></td><td align="left" valign="top"> J. Checa (DAOM 234345), Guanacaste, COSTA RICA
</td><td align="left" valign="top"> DQ402184
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"><italic>Od. sinense</italic></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123345&link_type=cbs">CBS 123345</ext-link></bold></td><td align="left" valign="top"> M. Gryzenhout (BPI 878730), Limpopo, SOUTH AFRICA
</td><td align="left" valign="top"> FJ161169
</td><td align="left" valign="top"> FJ161209
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> FJ161130
</td></tr><tr><td align="left" valign="top"></td><td align="left" valign="top"><bold>EB 0339</bold></td><td align="left" valign="top"> E.W.A. Boehm (BPI 879800), NJ, U.S.A.
</td><td align="left" valign="top"><bold>GU397364</bold></td><td align="left" valign="top"><bold>GU397348</bold></td><td align="left" valign="top"><bold>GU397339</bold></td><td align="left" valign="top"><bold>GU397357</bold></td></tr><tr><td align="left" valign="top"><italic>Opegrapha dolomitica</italic></td><td align="left" valign="top"><bold>DUKE 0047528</bold></td><td align="left" valign="top"> C. Gueidan (AFTOL 993), CROATIA
</td><td align="left" valign="top"> DQ883706
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> DQ883732
</td><td align="left" valign="top"> DQ883714
</td></tr><tr><td align="left" valign="top"><italic>Ostreichnion curtisii</italic></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=198.34&link_type=cbs">CBS 198.34</ext-link></bold></td><td align="left" valign="top"> M.L. Lohman (No. 464), GA, U.S.A.
</td><td align="left" valign="top"> FJ161137
</td><td align="left" valign="top"> FJ161176
</td><td align="left" valign="top"> FJ161093
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"><italic>O. sassafras</italic></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=322.34&link_type=cbs">CBS 322.34</ext-link></bold></td><td align="left" valign="top"> M.L. Lohman (No. 530), NC, U.S.A.
</td><td align="left" valign="top"> FJ161148
</td><td align="left" valign="top"> FJ161188
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> FJ161122
</td></tr><tr><td align="left" valign="top"><italic>Patellaria atrata</italic></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=958.97&link_type=cbs">CBS 958.97</ext-link></bold></td><td align="left" valign="top"> G. Marson, Wasserbillig, Bahnhof, LUXEMBOURG
</td><td align="left" valign="top"><bold>GU296181</bold></td><td align="left" valign="top"><bold>GU301855</bold></td><td align="left" valign="top"><bold>GU349038</bold></td><td align="left" valign="top"><bold>GU371726</bold></td></tr><tr><td align="left" valign="top"><italic>Phoma herbarum</italic></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=276.37&link_type=cbs">CBS 276.37</ext-link></bold></td><td align="left" valign="top"> AFTOL 1575
</td><td align="left" valign="top"> DQ678014
</td><td align="left" valign="top"> DQ678066
</td><td align="left" valign="top"> DQ677909
</td><td align="left" valign="top"> DQ677962
</td></tr><tr><td align="left" valign="top"><italic>Pleospora herbarum</italic></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=191.86&link_type=cbs">CBS 191.86</ext-link></bold></td><td align="left" valign="top"> E.G. Simmons, AFTOL_940, Uttar Pradesh, INDIA
</td><td align="left" valign="top"> DQ247812
</td><td align="left" valign="top"> DQ247804
</td><td align="left" valign="top"> DQ471090
</td><td align="left" valign="top"> DQ247794
</td></tr><tr><td align="left" valign="top"><italic>Psiloglonium araucanum</italic></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=112412&link_type=cbs">CBS 112412</ext-link></bold></td><td align="left" valign="top"> S. Marincowitz (PREM 57570), Kirstenbosch, SOUTH AFRICA
</td><td align="left" valign="top"> FJ161133
</td><td align="left" valign="top"> FJ161172
</td><td align="left" valign="top"> FJ161089
</td><td align="left" valign="top"> FJ161112
</td></tr><tr><td align="left" valign="top"></td><td align="left" valign="top"><bold>CMW 18760</bold></td><td align="left" valign="top"> S. Marincowitz (PREM 57569), Kirstenbosch, SOUTH AFRICA
</td><td align="left" valign="top"> FJ161151
</td><td align="left" valign="top"> FJ161192
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"></td><td align="left" valign="top"><bold>CMW 17941</bold></td><td align="left" valign="top"> S. Marincowitz (PREM 57566), Jonkershoek, SOUTH AFRICA
</td><td align="left" valign="top"> FJ161149
</td><td align="left" valign="top"> FJ161190
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"><italic>P. clavisporum</italic></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123338&link_type=cbs">CBS 123338</ext-link></bold></td><td align="left" valign="top"> E.W.A. Boehm (BPI 878726), NJ, U.S.A.
</td><td align="left" valign="top"> FJ161156
</td><td align="left" valign="top"> FJ161197
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> FJ161123
</td></tr><tr><td align="left" valign="top"></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123339&link_type=cbs">CBS 123339</ext-link></bold></td><td align="left" valign="top"> E.W.A. Boehm (BPI 878727), NJ, U.S.A.
</td><td align="left" valign="top"> FJ161157
</td><td align="left" valign="top"> FJ167526
</td><td align="left" valign="top"> FJ161105
</td><td align="left" valign="top"> FJ161124
</td></tr><tr><td align="left" valign="top"></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123340&link_type=cbs">CBS 123340</ext-link></bold></td><td align="left" valign="top"> E.W.A. Boehm (BPI 878728), NJ, U.S.A.
</td><td align="left" valign="top"> FJ161165
</td><td align="left" valign="top"> FJ161205
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123341&link_type=cbs">CBS 123341</ext-link></bold></td><td align="left" valign="top"> E.W.A. Boehm (BPI 878729), NJ, U.S.A.
</td><td align="left" valign="top"> FJ161166
</td><td align="left" valign="top"> FJ161206
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"></td><td align="left" valign="top"><bold>GKM 344A</bold></td><td align="left" valign="top"> G.K. Mugambi (BPI 879801, EA), Malindi, KENYA
</td><td align="left" valign="top"><bold>GU397365</bold></td><td align="left" valign="top"> GQ221889
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"></td><td align="left" valign="top"><bold>GKM L172A</bold></td><td align="left" valign="top"> G.K. Mugambi (EA), Malindi, KENYA
</td><td align="left" valign="top"><bold>GU323192</bold></td><td align="left" valign="top"><bold>GU323204</bold></td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"><italic>P. simulans</italic></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=206.34&link_type=cbs">CBS 206.34</ext-link></bold></td><td align="left" valign="top"> M.L. Lohman, MI, U.S.A.
</td><td align="left" valign="top"> FJ161139
</td><td align="left" valign="top"> FJ161178
</td><td align="left" valign="top"> FJ161094
</td><td align="left" valign="top"> FJ161116
</td></tr><tr><td align="left" valign="top"></td><td align="left" valign="top"><bold>ANM 1557</bold></td><td align="left" valign="top"> A.N. Miller (BPI 879803, ILLS), Smoky Mts., TN, U.S.A.
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> GQ221873
</td><td align="left" valign="top"> GQ221920
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"><italic>Quasiconcha reticulata</italic></td><td align="left" valign="top"><bold>EB QR</bold></td><td align="left" valign="top"> M. Blackwell (RLG 14189), AZ, U.S.A.
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"><bold>GU397349</bold></td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"><italic>Roccella fuciformis</italic></td><td align="left" valign="top"><bold>AFTOL 126</bold></td><td align="left" valign="top"> Diederich 15572
</td><td align="left" valign="top"> AY584678
</td><td align="left" valign="top"> AY584654
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> DQ782866
</td></tr><tr><td align="left" valign="top"><italic>Rhytidhysteron hysterinum</italic></td><td align="left" valign="top"><bold>EB 0351</bold></td><td align="left" valign="top"> A. Gardiennet (BPI 879804), Gevrey-Chambertin, FRANCE
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"><bold>GU397350</bold></td><td align="left" valign="top"><bold>GU397340</bold></td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"><italic>R. opuntiae</italic></td><td align="left" valign="top"><bold>GKM 1190</bold></td><td align="left" valign="top"> G.K. Mugambi (BPI 879805, EA), Malindi, KENYA
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"> GQ221892
</td><td align="left" valign="top"><bold>GU397341</bold></td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"><italic>R. rufulum</italic></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=306.38&link_type=cbs">CBS 306.38</ext-link></bold></td><td align="left" valign="top"> R.K. Voorhees (AFTOL 2109), EUROPE
</td><td align="left" valign="top"> AF164375
</td><td align="left" valign="top"> FJ469672
</td><td align="left" valign="top"><bold>GU349031</bold></td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"></td><td align="left" valign="top"><bold>GKM 361A</bold></td><td align="left" valign="top"> G.K. Mugambi (BPI 879806, EA), Malindi, KENYA
</td><td align="left" valign="top"><bold>GU296192</bold></td><td align="left" valign="top"> GQ221893
</td><td align="left" valign="top"><bold>GU349031</bold></td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"></td><td align="left" valign="top"><bold>EB 0381</bold></td><td align="left" valign="top"> E. Nkansah (BPI 879807), Kwame Nkrumah, GHANA
</td><td align="left" valign="top"><bold>GU397366</bold></td><td align="left" valign="top"><bold>GU397351</bold></td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"></td><td align="left" valign="top"><bold>EB 0382</bold></td><td align="left" valign="top"> E. Nkansah (BPI 879808), Kwame Nkrumah, GHANA
</td><td align="left" valign="top"> —
</td><td align="left" valign="top"><bold>GU397352</bold></td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"></td><td align="left" valign="top"><bold>EB 0383</bold></td><td align="left" valign="top"> E. Nkansah (BPI 879809), Kwame Nkrumah, GHANA
</td><td align="left" valign="top"><bold>GU397353</bold></td><td align="left" valign="top"><bold>GU397367</bold></td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"></td><td align="left" valign="top"><bold>EB 0384</bold></td><td align="left" valign="top"> E. Nkansah (BPI 879810), Kwame Nkrumah, GHANA
</td><td align="left" valign="top"><bold>GU397368</bold></td><td align="left" valign="top"><bold>GU397354</bold></td><td align="left" valign="top"> —
</td><td align="left" valign="top"> —
</td></tr><tr><td align="left" valign="top"><italic>Scorias spongiosa</italic></td><td align="left" valign="top"><bold><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=325.33&link_type=cbs">CBS 325.33</ext-link></bold></td><td align="left" valign="top"> L.H. Leonian (AFTOL 1594)
</td><td align="left" valign="top"> DQ678024
</td><td align="left" valign="top"> DQ678075
</td><td align="left" valign="top"> DQ677920
</td><td align="left" valign="top"> DQ677973
</td></tr><tr><td align="left" valign="top"><italic>Simonyella variegata</italic></td><td align="left" valign="top"><bold>AFTOL 80</bold></td><td align="left" valign="top">DUKE Printzen 14310a
</td><td align="left" valign="top">AY584669
</td><td align="left" valign="top">—
</td><td align="left" valign="top">DQ782891
</td><td align="left" valign="top">DQ782861
</td></tr></tbody></table><table-wrap-foot><fn><p><bold>AFTOL:</bold> Assembling the Fungal Tree of Life; <bold>BPI:</bold> United States
USDA ARS National Fungus Collections, Beltsville, MD; <bold>CBS:</bold>Centraalbureau voor Schimmelcultures, Utrecht, The Netherlands; <bold>CMW:</bold>Forestry & Agricultural Biotechnology Institute (FABI), University of
Pretoria, Pretoria, Republic of South Africa; <bold>DAOM:</bold> National
Mycological Herbarium, Department of Agriculture, Ottawa, Ontario, Canada;
<bold>DUKE:</bold> Duke University Herbarium, Durham, North Carolina; <bold>EA:</bold>National Museums of Kenya East African Herbarium, Nairobi, Kenya; <bold>F:</bold>Field Museum of Natural History, Chicago, IL; <bold>ILLS:</bold> Illinois Natural
History Survey Herbarium, Champaign, IL; <bold>PREM:</bold> The South African
National Collection of Fungi, National Mycological Herbarium, Pretoria, South
Africa; <bold>RLG:</bold> The Robert L. Gilbertson Mycological Herbarium at the
University of Arizona. Culture and specimen abbreviations: ANM: A.N. Miller;
EB: E.W.A. Boehm; GKM: G.K. Mugambi, SMH: S.M. Huhndorf. GenBank accessions
marked in bold represent new sequences generated in the current study.</p></fn></table-wrap-foot></table-wrap></p><p>An attempt was made to include a broad range of fungal isolates, belonging
to or previously affiliated with the <italic>Hysteriaceae, Mytilinidiaceae,
Gloniaceae</italic> and <italic>Patellariaceae</italic>(<xref ref-type="table" rid="tbl1">Table 1</xref>). A geographically
diverse (Cuba, Europe, Ghana, Kenya, New Zealand, South Africa, Tasmania,
North and South America) and high density taxon sampling strategy was
employed. This included multiple isolates/species from the genera:
<italic>Anteaglonium</italic> (6/4), <italic>Encephalographa</italic> (1/1),
<italic>Farlowiella</italic> (3/1), <italic>Gloniopsis</italic> (8/4), <italic>Glonium</italic> (4/2),
<italic>Hysterium</italic> (12/5), <italic>Hysterobrevium</italic> (14/3),
<italic>Hysterographium</italic> (2/1), <italic>Hysteropatella</italic> (2/2),
<italic>Lophium</italic> (4/2), <italic>Mytilinidion</italic> (13/10), <italic>Oedohysterium</italic>(5/3), <italic>Ostreichnion</italic> (2/2), <italic>Patellaria</italic> (1/1),
<italic>Psiloglonium</italic> (11/3), <italic>Quasiconcha</italic> (1/1),
<italic>Rhytidhysteron</italic> (8/3), and 24 outgroup taxa, for a total of 121 taxa.
All cultures and the herbarium specimens from which they were derived, have
been deposited and are permanently conserved in the certified public
institutions given in <xref ref-type="table" rid="tbl1">Table
1</xref>.</p><p>Within the <italic>Pleosporales</italic>, we sampled <italic>Anteaglonium abbreviatum,
A. globosum, A. latirostrum</italic> and <italic>A. parvulum, Byssothecium circinans,
Cochliobolus heterostrophus, Delitschia winteri, Herpotrichia diffusa,
Leptosphaeria maculans, Phoma herbarum</italic>, and <italic>Pleospora herbarum</italic>.
Eight representatives from the <italic>Dothideomycetidae</italic> were included as
outgroups to the <italic>Pleosporomycetidae</italic>, namely <italic>Elsinoë
veneta</italic> and <italic>Myriangium duriaei</italic> (<italic>Myriangiales</italic>),
<italic>Dothidea sambuci</italic> and <italic>D. insculpta</italic> (<italic>Dothideales</italic>),
<italic>Mycosphaerella punctiformis</italic> and <italic>Scorias spongiosa</italic>(<italic>Capnodiales</italic>). <italic>Botryosphaeria dothidea</italic>, and <italic>Guignardia
gaultheriae</italic> (<italic>Botryosphaeriales</italic>). <italic>Jahnula aquatica</italic> and
<italic>Aliquandostipite khaoyaiensis</italic> (<italic>Jahnulales</italic>), were also
included. Four taxa in the <italic>Arthoniomycetes</italic>, were used as outgroups to
the <italic>Dothideomycetes</italic>, namely <italic>Opegrapha dolomitica, Simonyella
variegata, Roccella fuciformis</italic>, and <italic>Arthonia caesia</italic>. These are
not presented in <xref ref-type="fig" rid="fig1">Fig. 1</xref>, due to
space limitations, but are presented as a full tree available on TreeBASE, as
well as in <xref ref-type="table" rid="tbl1">Table 1</xref>.</p></sec><sec><title>DNA extraction, amplification and sequencing</title><p>Genomic DNA was recovered using the DNeasy® Plant Mini Kit (Qiagen
Inc., Valencia, CA, U.S.A.), following the instructions of the manufacturer,
but using sterile white quartz sand and a Kontes® battery-powered pestle
grinder in 1.5 mL microfuge tubes. The nuSSU was amplified and double-strand
sequenced using the primers NS1 and NS4
(<xref ref-type="bibr" rid="ref120">White <italic>et al.</italic> 1990</xref>),
while amplification of the nuLSU utilised the primers LROR
(<xref ref-type="bibr" rid="ref90">Rehner & Samuels 1994</xref>)
and LR7 (<xref ref-type="bibr" rid="ref118">Vilgalys & Hester
1990</xref>), in addition to the internal sequencing primers LR3R and
LR16 (<xref ref-type="bibr" rid="ref79">Moncalvo <italic>et al.</italic>1993</xref>). Final concentrations for 50 μL PCR amplification
reactions were as follows: 1 μM of each forward and reverse primer, 2 mM
MgCl<sub>2</sub>, 200 μM dNTP, 1X Promega GoTaq® Flexi Reaction Buffer,
1.25 U of Promega GoTaq® Polymerase, and 2 μL template DNA diluted
tenfold. For the nuSSU and nuLSU, PCR reaction parameters were as follows: a
95 °C pre-melt for 3 min, and 35 cycles of 95 °C for 20 s, 54 °C
for 30 s and 72 °C for 60 s, followed by a final extension at 72 °C
for 10 min. For <italic>TEF1</italic> and <italic>RPB2</italic>, PCR amplification conditions
followed those in Schoch <italic>et al.</italic>(<xref ref-type="bibr" rid="ref100">2006</xref>). Primers used for the
amplifications and sequencing of these protein coding genes were for
<italic>TEF1</italic>: 983 & 2218R; and for <italic>RPB2</italic>: fRPB2-5F &
fRPB2-7cR. PCR reactions were performed using PCR Master Mix Polymerase from
Promega Corporation (Fitchburg, Wisconsin, U.S.A.) and run on an iCycler®
from Biorad (Hercules, California, U.S.A.). For the amplification of DNA
fragments used to infer the <italic>TEF1</italic> amino acid sequence, the following
conditions were used: (1) 94 °C for 2 min; (2) five cycles of 94 °C
for 40 s, 55 °C for 45 s lowering by 0.8 °C per cycle and 72 °C
for 90 s; (3) 30 cycles of 94 °C for 30 s, 52 °C for 45 s and 72
°C for 120 s and (4) a cycle for 10 min at 72 °C. Amplifications of
DNA fragments used to infer the <italic>RPB2</italic> amino acid sequence utilised the
same cycle parameters, except for changes in steps (2) and (3) where the
annealing temperatures of 55 °C and 52 °C were changed to 50 °C
and 45 °C, respectively. Amplified PCR products were cleaned using the
QIAquick® PCR Purification Kit (Qiagen Inc.) and resuspended in water
prior to outsourcing for sequencing (Macrogen U.S.A., Inc.).</p><p><fig position="float" id="fig1"><label>Fig. 1.</label><caption><p>Combined ribosomal (nuSSU & nuLSU) and protein coding gene
(<italic>TEF1</italic> & <italic>RPB2</italic>) DNA phylogeny for bitunicate ascomycetes
belonging to or previously affiliated with the <italic>Hysteriaceae</italic>,
<italic>Mytilinidiaceae</italic>, <italic>Gloniaceae</italic> and <italic>Patellariaceae</italic>.
Also included are representatives from allied groups such as the
<italic>Pleosporales</italic>, <italic>Jahnulales</italic>, <italic>Patellariales</italic>, and
<italic>Botryosphaeriales</italic>, as well as representatives from the
<italic>Dothideales</italic>, <italic>Myriangiales</italic> and <italic>Capnodiales</italic> in the
<italic>Dothideomycetidae</italic>. The <italic>Arthoniomycetes</italic>, chosen as outgroup,
are not presented here due to space limitations, but are available in the full
tree on TreeBASE. The tree is the highest scoring tree obtained by maximum
likelihood in RAxML. Nodal values, given as percentages, are as follows:
Bayesian posterior probability / maximum likelihood bootstrap. Only values
above 50 % are shown.</p></caption><graphic xlink:href="49fig1"></graphic></fig></p></sec><sec><title>Phylogenetic analysis</title><p>DNA sequences were derived from previous studies
(<xref ref-type="bibr" rid="ref100">Schoch <italic>et al.</italic> 2006</xref>,
<xref ref-type="bibr" rid="ref18">Boehm <italic>et al.</italic> 2009</xref>,
<xref ref-type="bibr" rid="ref81">Mugambi & Huhndorf 2009</xref>),
as well as from a number of new accessions generated in this study
(<xref ref-type="table" rid="tbl1">Table 1</xref>). Sequences were
aligned using default options for a simultaneous method of estimating
alignments and tree phylogenies, SATé
(<xref ref-type="bibr" rid="ref53">Liu <italic>et al.</italic> 2009</xref>).
Protein coding fragments were translated using BioEdit v. 7.0.1
(<xref ref-type="bibr" rid="ref39">Hall 2004</xref>), and aligned
within SATé as amino acid sequence data. These were then aligned with
their respective DNA sequences using the RevTrans v. 1.4 Server
(<xref ref-type="bibr" rid="ref119">Wernersson & Pedersen
2003</xref>). Newly generated sequences were subsequently added to the
core alignment with MAFFT v. 6.713 (<xref ref-type="bibr" rid="ref44">Katoh
<italic>et al.</italic> 2009</xref>). A supermatrix of four genes (nuLSU, nuSSU
<italic>TEF1, RPB2</italic>) consisting of 56 % gaps and undetermined characters,
across 121 taxa was obtained.</p><p>The matrix was analysed using maximum likelihood in RAxML v. 7.0.4
(<xref ref-type="bibr" rid="ref107">Stamatakis 2006</xref>). Data was
partitioned by individual gene and, where applicable, by codon, as in Schoch
<italic>et al.</italic> (2009). A most likely tree was obtained after 100 successive
searches in RAxML under the GTR model with gamma rate distribution across 11
partitions and starting each search from a randomised tree with a rapid hill
climbing option and joint branch length optimisation. Five hundred fast
bootstrap pseudoreplicates (<xref ref-type="bibr" rid="ref108">Stamatakis
<italic>et al.</italic> 2008</xref>) were run under the same conditions and these
values are given above each node. The matrix analysed in this study produced
4174 distinct alignment patterns and the most likely tree had a log likelihood
of -72114.22899. The average log likelihood over 100 trees was -72117.730727.
Three independent Bayesian phylogenetic analyses were performed in MrBayes
3.1.2 (<xref ref-type="bibr" rid="ref42">Huelsenbeck & Ronquist
2001</xref>) using a uniform [GTR+I+G] model. The Metropolis-coupled
Markov chain Monte Carlo (MCMC) sampling approach was used to calculate
posterior probabilities (PP). For each Bayesian run four Markov chains were
run from a random starting tree for 5 000 000 generations and trees sampled
every 100 generations. The first 50 000 generation trees were discarded as
burn-in prior to convergence of four of the chains. All three runs reached a
plateau that converged. One run was chosen to construct a 50 % majority rule
consensus tree of all trees remaining after the burn in was discarded.
Bayesian Posterior Probabilities with those equal or greater than 50 % are
given below each node (<xref ref-type="fig" rid="fig1">Fig.
1</xref>).</p></sec></sec><sec><title>RESULTS AND DISCUSSION</title><sec><title>Phylogenetic analysis – ordinal level</title><p>At the ordinal level in the <italic>Pleosporomycetidae</italic>, molecular data
indicate that the <italic>Hysteriales</italic> are closely related to the
<italic>Pleosporales</italic> (<xref ref-type="fig" rid="fig1">Fig. 1</xref>),
as was indicated in earlier studies
(<xref ref-type="bibr" rid="ref100">Schoch <italic>et al.</italic> 2006</xref>,
<xref ref-type="bibr" rid="ref18">Boehm <italic>et al.</italic> 2009</xref>).
This is also confirmed by morphological evidence related to the centrum. Thus,
the <italic>Hysteriales</italic> share a very similar centrum with the
<italic>Pleosporales</italic>, that is, one defined by the <italic>Pleospora</italic>-type, in
which cellular, septate pseudoparaphyses grow downwards from the cavity roof,
initially anchored at both ends, and occupy the locule prior to the formation
of asci (<xref ref-type="bibr" rid="ref67">Luttrell 1951</xref>).
However, there is also strong branch support for its separation from the
<italic>Pleosporales</italic> (<xref ref-type="bibr" rid="ref18">Boehm <italic>et
al.</italic> 2009</xref>). The <italic>Hysteriales</italic> are therefore retained as
defined by Luttrell (<xref ref-type="bibr" rid="ref69">1955</xref>), to
emphasise the elongated hysteriaceous locule, capable of relatively
indeterminate linear growth, as distinct from the strict
<italic>Pleospora</italic>-type centrum, defined as it is by constrained concentric
growth. In contrast to the close association between the <italic>Hysteriales</italic>and the <italic>Pleosporales</italic>, the <italic>Mytilinidiales</italic> forms a more
distant clade within the <italic>Pleosporomycetidae</italic>(<xref ref-type="bibr" rid="ref18">Boehm <italic>et al.</italic>2009</xref>).</p></sec><sec><title>Phylogenetic analysis – family level</title><sec><title>Hysteriaceae</title><p>Although the <italic>Hysteriales</italic> receives high branch support as a
monophyletic entity, distinct from the closely related <italic>Pleosporales</italic>,
two major groups can be defined within the family. The first supports Clades
A–C, whereas the second supports Clades D and E
(<xref ref-type="fig" rid="fig1">Fig. 1</xref>).</p><p><bold>Clade A:</bold> This first clade is characterised by <italic>Hysterographium
mori</italic>, with short pigmented dictyospores, <italic>Gloniopsis constricta</italic>,
and <italic>Gp. smilacis</italic>, the latter two with short hyaline dictyospores. The
<italic>Gp. smilacis</italic> isolates originate from highly diverse geographical
sources (<italic>e.g.</italic>, Sweden, South Africa, North America;
<xref ref-type="table" rid="tbl1">Table 1</xref>), thus strongly
supporting its phylogenetic placement. As these taxa are far removed from the
types for their respective genera, we propose here to unite them in
<italic>Hysterobrevium gen. nov.</italic>, as <italic>Hb. mori comb. nov., Hb. constrictum
comb. nov.</italic>, and <italic>Hb. smilacis comb. nov.</italic></p><p><bold>Clade B:</bold> This clade (<xref ref-type="fig" rid="fig1">Fig.
1</xref>) appears monophyletic for the newly reinstated genus
<italic>Psiloglonium</italic> (<xref ref-type="bibr" rid="ref18">Boehm <italic>et
al.</italic> 2009</xref>), with hyaline didymospores. It includes the
following species: <italic>P. simulans</italic>, <italic>P. clavisporum,</italic> and <italic>P.
araucanum comb. nov.</italic> In this study, we propose a number of new
combinations for the genus <italic>Psiloglonium</italic>, with <italic>P. lineare</italic> as
the type (<xref ref-type="bibr" rid="ref18">Boehm <italic>et al.</italic>2009</xref>), to accommodate species previously classified under the
genus <italic>Glonium</italic>, now in the <italic>Gloniaceae</italic>.</p><p><bold>Clade C:</bold> This clade is characterised by pigmented phragmospores
belonging to four species of the genus <italic>Hysterium</italic>, namely <italic>H.
pulicare, H. angustatum, H. vermiforme</italic>, which have 3-septate spores, and
<italic>H. barrianum sp. nov.</italic>, which has 9-septate spores. Again, a
geographically diverse set of isolates were surveyed
(<xref ref-type="table" rid="tbl1">Table 1</xref>). For instance, taxon
sampling for <italic>H. angustatum</italic> included isolates originating from Kenya,
New Zealand, South Africa, and North America
(<xref ref-type="fig" rid="fig1">Fig. 1</xref>). Also within this
clade, but with weak bootstrap support, is <italic>Ostreichnion sassafras</italic>,
and <italic>O. curtisii</italic>, previously transferred from the
<italic>Mytilinidiaceae</italic> to the <italic>Hysteriaceae</italic>(<xref ref-type="bibr" rid="ref18">Boehm <italic>et al.</italic>2009</xref>).</p><p><bold>Clade D:</bold> This clade is heterogeneous, but can be divided into two
sub-clades. The first sub-clade includes two species formerly in the genus
<italic>Hysterium</italic>, namely <italic>H. insidens</italic> and <italic>H. sinense</italic>.
Molecular data indicate that these species are not related to the type
species, <italic>H. pulicare</italic>, nor to related species within Clade C.
Morphology also supports this separation, as <italic>H. insidens</italic> and <italic>H.
sinense</italic> both possess phragmospores with a swollen supra-median cell. We
therefore propose <italic>Oedohysterium gen. nov.</italic>, to accommodate <italic>Od.
insidens comb. nov.</italic> and <italic>Od. sinense comb. nov.</italic> Also grouping in
Clade D is <italic>Hysterographium pulchrum</italic>. Despite the fact that <italic>Hg.
pulchrum</italic> possesses dictyospores, we propose to unite it within
<italic>Oedohysterium</italic>, as <italic>Od. pulchrum comb. nov.</italic>, on account that
it too possesses a swollen supra-median cell. Also present in this subclade
are two isolates of <italic>Hb. mori</italic>, distant from the other <italic>Hb.
mori</italic> accessions in Clade A; this anomaly will be discussed later. A
separate subclade is evident in Clade D, and defines the type species for the
genus <italic>Gloniopsis</italic>, namely <italic>Gp. praelonga</italic>. Closely associated
with <italic>Gp. praelonga</italic> is one representative of <italic>Hg. subrugosum</italic>.
Dictyospores of both species are of similar shape, size and degree of
septation, differing only in the lack of pigmentation and a gelatinous sheath.
We thus propose that <italic>Gp. praelonga</italic> and <italic>Hg. subrugosum</italic> be
united within the same genus, proposing <italic>Gloniopsis subrugosa comb.
nov</italic>. The other two representatives of <italic>Gp. subrugosa</italic> do not fall
into Clade D, but lie adjacent. Lastly, an additional two species are
described in Clade D, namely <italic>Gloniopsis arciformis sp. nov.</italic> and
<italic>Gp. kenyensis sp. nov.</italic>, both from East Africa
(<xref ref-type="table" rid="tbl1">Table 1</xref>).</p><p><bold>Clade E:</bold> This clade is well-supported and defines two species in the
genus <italic>Rhytidhysteron</italic>, namely <italic>R. rufulum</italic>, and <italic>R.
hysterinum</italic>. Taxon sampling included isolates originating from France,
Ghana, Kenya and North America. This clade therefore supports the transference
of the genus <italic>Rhytidhysteron</italic> from the <italic>Patellariaceae</italic> to the
<italic>Hysteriaceae</italic>, as initially proposed by Boehm <italic>et al.</italic>(<xref ref-type="bibr" rid="ref18">2009</xref>). The third species of
<italic>Rhytidhysteron, R. opuntiae</italic>, is distant to the first two species, but
remains adjacent to Clade E.</p></sec><sec><title>Mytilinidiaceae</title><p>In contrast to the <italic>Hysteriales</italic>, the family
<italic>Mytilinidiaceae</italic> represents a highly monophyletic entity, defining the
order <italic>Mytilinidiales</italic> (<xref ref-type="bibr" rid="ref18">Boehm
<italic>et al.</italic> 2009</xref>). The conchate nature of the fruitbody and
the thin-walled peridium, seem to unite what at first may seem a disparate
group of fungi into a single family (<xref ref-type="fig" rid="fig1">Fig.
1</xref>). In this study, we have sampled 10 of the 15 species of
<italic>Mytilinidion</italic>, characterised by phragmospores and scolecospores, two
of the four species of <italic>Lophium</italic>, with filiform spores, as well as the
monotypic <italic>Quasiconcha</italic>, with reticulated 1-septate spores
(<xref ref-type="table" rid="tbl1">Table 1</xref>). Although
monophyletic, sequence data also indicate a complex pattern of speciation
within the family, one that is not premised on past assumptions based on spore
morphology (<xref ref-type="fig" rid="fig1">Fig. 1</xref>).</p></sec><sec><title>Gloniaceae</title><p>As for the monotypic family <italic>Gloniaceae</italic>(<xref ref-type="bibr" rid="ref18">Boehm <italic>et al.</italic> 2009</xref>),
based on the genus <italic>Glonium</italic>, previously classified within the
<italic>Hysteriaceae</italic> (<xref ref-type="bibr" rid="ref122">Zogg
1962</xref>), surprisingly, sequence data indicate that it finds close
affinity with the <italic>Mytilinidiaceae</italic>(<xref ref-type="fig" rid="fig1">Fig. 1</xref>). This is based on four
isolates, representing two species, <italic>Glonium stellatum</italic> and <italic>G.
circumserpens</italic>. However, the <italic>Gloniaceae</italic> is not included within
the <italic>Mytilinidiales</italic>, due to the highly divergent morphology associated
with the genus <italic>Glonium</italic>. These include character states related to the
hamathecium (persistent cellular pseudoparaphyses <italic>versus</italic> narrow
trabeculate pseudoparaphyses) and to the fruitbody (dichotomously branched
<italic>versus</italic> conchate), for the <italic>Gloniaceae</italic> and
<italic>Mytilinidiaceae</italic>, respectively. Thus, for the present, we propose that
the family <italic>Gloniaceae</italic> be considered <italic>Pleosporomycetidae fam.
incertae sedis</italic>.</p></sec></sec></sec><sec><title>TAXONOMY</title><p><italic><bold>Hysteriaceae</bold></italic> Chevall. 1826, <italic><bold>Hysteriales</bold></italic>Lindau 1897.</p><p>Fungi classified in the <italic>Hysteriaceae</italic>(<xref ref-type="bibr" rid="ref22">Chevallier 1826</xref>) have been
traditionally defined by a specialised ascocarp termed the hysterothecium
(<xref ref-type="bibr" rid="ref24">Clements 1909</xref>). Hysterothecia
are dense, persistent carbonaceous structures, distinctly navicular in
outline, and bear a pronounced longitudinal slit running the length of the
long axis of the fruitbody. They can be immersed to erumpent to entirely
superficial, solitary to gregarious, ellipsoid to greatly elongated, sometimes
branched or triradiate. In vertical section, hysterothecia are globose to
obovoid, typically with a thick three-layered peridium, composed of small
pseudoparenchymatous cells, the outer layer heavily encrusted with pigment and
often longitudinally striated on the surface, the middle layer lighter in
pigmentation and the inner layer distinctly thin-walled, pallid and compressed
(<xref ref-type="bibr" rid="ref7">Barr 1987</xref>). The hamathecium is
composed of persistent, narrow cellular pseudoparaphyses, often borne in a gel
matrix, with tips darkened or branched at maturity above the asci. Bitunicate
asci are borne in a basal layer and at maturity are typically clavate to
cylindric, bearing 8 ascospores, overlapping biseriate, ranging from hyaline
to dark brown, obovoid, clavate, ellipsoid or fusoid. Ascospores are highly
diverse in septation and range from didymospores to phragmospores to
dictyospores, at times surrounded by a gel coating, and often show bipolar
asymmetry (<xref ref-type="bibr" rid="ref7">Barr 1987</xref>). Zogg
(<xref ref-type="bibr" rid="ref122">1962</xref>) accepted the following
seven genera within the <italic>Hysteriaceae</italic>: <italic>Farlowiella, Gloniella,
Gloniopsis, Glonium, Hysterium, Hysterocarina</italic>, and
<italic>Hysterographium</italic>.</p><p>The traditional circumspection of the <italic>Hysteriaceae</italic> was based on
character states related to the hysterothecium and spore morphology
(<italic>e.g.</italic>, septation and pigmentation), character states previously
considered synapomorphic (<xref ref-type="bibr" rid="ref122">Zogg
1962</xref>). However, recent molecular data underscore the potential for
morphology to be difficult to interpret, and even unhelpful in phylogenetic
inference and reconstruction for this group of fungi
(<xref ref-type="bibr" rid="ref100">Schoch <italic>et al.</italic> 2006</xref>,
<xref ref-type="bibr" rid="ref18">Boehm <italic>et al.</italic> 2009</xref>,
<xref ref-type="bibr" rid="ref81">Mugambi & Huhndorf 2009</xref>).
Thus, a number of examples of convergent evolution are presented in the
current study, which relate not only to the fruitbody, but to spore morphology
as well. As a result, three genera have been removed from the family
(<italic>Glonium, Hysterographium</italic> and <italic>Farlowiella</italic>), based on
convergence associated with the fruitbody. Additionally, within the family,
several genera have their members spanning different clades
(<xref ref-type="fig" rid="fig1">Fig. 1</xref>). This necessitated the
description of two new genera (<italic>Oedohysterium</italic> and
<italic>Hysterobrevium</italic>), as well as three new species, one in
<italic>Hysterium</italic> and two in <italic>Gloniopsis</italic>, in addition to a number of
new combinations involving <italic>Psiloglonium, Oedohysterium,
Hysterobrevium</italic> and <italic>Gloniopsis</italic>. These taxonomic changes have
de-emphasised both spore septation and spore pigmentation as reliable
character states for deducing phylogenetic relationships within the family.
Nevertheless, in the keys that follow, we have endeavoured to provide a
morphological basis for the new phylogenies revealed by molecular data.</p><p>Data have also necessitated that we expand the concept of the
<italic>Hysteriaceae</italic> to include thin-walled mytilinidioid forms previously in
the <italic>Mytilinidiaceae</italic> (<italic>e.g., Ostreichnion</italic>), as well as
patellarioid forms previously in the <italic>Patellariaceae</italic> (<italic>e.g.,
Rhytidhysteron</italic>). The inclusion of <italic>Ostreichnion</italic> within the
<italic>Hysteriaceae</italic> was unexpected. Unlike most members of the family, the
peridium in <italic>Ostreichnion</italic> is sclerenchymatoid and thin-walled,
defining a fragile mytilinidioid ascoma, and with a hamathecium typified by
trabeculate pseudoparaphyses (Barr
<xref ref-type="bibr" rid="ref4">1975</xref>,
<xref ref-type="bibr" rid="ref8">1990a</xref>). Including the genus
<italic>Ostreichnion</italic> in the <italic>Hysteriaceae</italic> implies that, either
morphological features within the genus need to be re-evaluated, or that the
family <italic>Hysteriaceae</italic> must also encompass mytilinidioid forms. More
difficult to understand perhaps is the inclusion of the genus
<italic>Rhytidhysteron</italic> within the <italic>Hysteriaceae</italic>. Although included
within the <italic>Patellariaceae</italic>(<xref ref-type="bibr" rid="ref48">Kutorga & Hawksworth
1997</xref>), phylogenetic data presented here and elsewhere
(<xref ref-type="bibr" rid="ref18">Boehm e<italic>t al.</italic> 2009</xref>),
clearly indicate that this genus lies quite distant from other members of the
<italic>Patellariaceae</italic>.</p><p>Some authors have included a number of additional genera within the
<italic>Hysteriaceae</italic>. For instance, the genera <italic>Hysteropatella,
Hysteroglonium</italic>, and <italic>Pseudoscypha</italic> were included in the
<italic>Hysteriaceae</italic> by Eriksson
(<xref ref-type="bibr" rid="ref33">2006</xref>). In addition, the
genera <italic>Hemigrapha, Graphyllium</italic>, and <italic>Encephalographa</italic> were
included in the family by Kirk <italic>et al.</italic>(<xref ref-type="bibr" rid="ref45">2001</xref>). In Boehm <italic>et
al.</italic> (<xref ref-type="bibr" rid="ref18">2009</xref>), two species
belonging to <italic>Hysteropatella</italic>, namely <italic>Hp. clavispora</italic>(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=247.34&link_type=cbs">CBS 247.34</ext-link>) and
<italic>Hp. elliptica</italic> (<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=935.97&link_type=cbs">CBS
935.97</ext-link>), did not cluster with any of the hysteriaceous taxa
surveyed. Instead, they formed a distant clade within the
<italic>Pleosporomycetidae</italic>, postulated to represent the emergence of the
<italic>Patellariales</italic>. In the present study, these two species of
<italic>Hysteropatella</italic> continue to be distant from the <italic>Hysteriaceae</italic>,
and also cluster now with <italic>Patellaria atrata</italic>(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=958.97&link_type=cbs">CBS 958.97</ext-link>).
Therefore, we do not include the genus <italic>Hysteropatella</italic> within the
<italic>Hysteriaceae</italic>.</p><p>Reid & Pirozynski
(<xref ref-type="bibr" rid="ref91">1966</xref>) in describing
<italic>Pseudoscypha abietis</italic> on the needles of <italic>Abies balsamea</italic> did
not mention the <italic>Hysteriaceae</italic>, and in fact stated that the fungus
cannot be assigned to any presently known order. In their illustrations, no
sterile tissue or excipulum is presented, and the bitunicate asci and
pseudoparaphyses arise directly from an erumpent orange basal stromatic
cushion. As such, we do not include <italic>Pseudoscypha</italic> as a member of the
<italic>Hysteriaceae</italic>. As for the genus <italic>Hemigrapha</italic>, Diederich &
Wedin (<xref ref-type="bibr" rid="ref30">2000</xref>) make the argument
for the inclusion of the genus in the <italic>Microthyriaceae</italic>, not the
<italic>Hysteriaceae</italic>. The genus <italic>Graphyllium</italic> possesses applanate,
clathrate ascospores borne in thin-walled membranous hysterothecia, at first
subcuticular, later erumpent, often associated with aquatic poaceous hosts.
The genus was included in the <italic>Hysteriaceae</italic> by Shoemaker & Babcock
(<xref ref-type="bibr" rid="ref103">1992</xref>) and Kirk <italic>et
al.</italic> (<xref ref-type="bibr" rid="ref45">2001</xref>), but was
earlier classified in the <italic>Phaeosphaeriaceae</italic>(<xref ref-type="bibr" rid="ref7">Barr 1987</xref>). A new species was
recently described from Costa Rica (<xref ref-type="bibr" rid="ref21">Checa
<italic>et al.</italic> 2007</xref>). The unique ascospore and the lack of
carbonisation or peridial wall thickness argue against the inclusion in the
<italic>Hysteriaceae</italic>, but molecular data are lacking.</p><sec><title>Key to the genera and allied genera of the <italic>Hysteriaceae</italic></title><p><list list-type="simple"><list-item><p>1. Ascomata apothecioid, opening widely when hydrated, fully exposing the
hymenium, which may be red or black (<italic>i.e.</italic>,
patellarioid)..............................................................................................................................................................
<italic><bold>Rhytidhysteron</bold></italic>1. Hysterothecia usually remaining closed,
or only opening slightly through a longitudinal fissure or sulcus to reveal a
lenticular, disk-like hymenium when hydrated and
mature....................................................................................................................
2</p></list-item><list-item><p>2. Ascospores pedicellate amerospores, the upper cell pigmented and much
larger than the lower, which remains un- or less-pigmented; anamorph
<italic>Acrogenospora</italic>.........................................................................................................................
<italic><bold>Farlowiella</bold>Note</italic>: The genus <italic>Farlowiella</italic> has been
removed from the <italic>Hysteriaceae</italic> and is currently listed as
<italic>Pleosporomycetidae gen. incertae sedis</italic>(<xref ref-type="bibr" rid="ref18">Boehm <italic>et al.</italic>2009</xref>).
2. Ascospores not as above, didymospores, phragmospores
or dictyospores, sometimes
pigmented........................................................ 3</p></list-item><list-item><p>3. Didymospores small, the two cells more or less equal in
size..................................................................................................................
4
3. Ascospores not as above, phragmospores, dictyospores, +/-
pigmentation, or very large didymospores (<italic>O.
curtisii</italic>)............................. 7</p></list-item><list-item><p>4. Ascospores
hyaline...................................................................................................................................................................................
5
4. Ascospores
pigmented..................................................................................................................................................
<italic><bold>Actidiographium</bold></italic></p></list-item><list-item><p>5. Didymospores less than 8 μm
long.....................................................................................................................................
<italic><bold>Anteaglonium</bold>Note</italic>: The genus <italic>Anteaglonium</italic> lies
within the <italic>Pleosporales</italic> (<xref ref-type="bibr" rid="ref81">Mugambi
& Huhndorf 2009</xref>), but is keyed out here with
<italic>Psiloglonium</italic>.
5. Didymospores longer than 8
μm...............................................................................................................................................................
6</p></list-item><list-item><p>6. Didymospores hyaline, borne in solitary or gregarious hysterothecia,
rarely associated with a subiculum, not laterally anastomosed to form
radiating stellate
composites.............................................................................................................
<italic><bold>Psiloglonium</bold>Note</italic>: One species of <italic>Anteaglonium, A.
latirostrum</italic>, will key out here, but belongs in the <italic>Pleosporales</italic>(<xref ref-type="bibr" rid="ref81">Mugambi & Huhndorf 2009</xref>)
and is also keyed out in the <italic>Psiloglonium</italic> key.
6. Didymospores
hyaline, borne in modified hysterothecia, usually associated with a subiculum,
strongly laterally anastomosed along their length to form radiating stellate
composites........................................................................................
<italic><bold>Glonium</bold>Note</italic>: The genus <italic>Glonium</italic> has been
transferred from the <italic>Hysteriaceae</italic> to the <italic>Gloniaceae</italic>,
currently listed as <italic>fam. incertae sedis</italic> within the
<italic>Pleosporomycetidae</italic> (<xref ref-type="bibr" rid="ref18">Boehm <italic>et
al.</italic> 2009</xref>).</p></list-item><list-item><p>7. Ascospores transversely septate phragmospores, or if with dictyospores
then also with red pigmentation.............................................
8
7. Ascospores transversely and longitudinally septate dictyospores, or
very large didymospores (<italic>O.
curtisii</italic>).......................................... 10</p></list-item><list-item><p>8. Ascospores hyaline
phragmospores............................................................................................................................................
<italic><bold>Gloniella</bold></italic>8. Ascospores pigmented phragmospores or in one
case (<italic>Od. pulchrum</italic>) with pigmented dictyospores and red pigmentation
in the
hamathecium.............................................................................................................................................................
9</p></list-item><list-item><p>9. Phragmospores 3-septate or rarely more, but without swollen supra-median
cell(s)...............................................................
<italic><bold>Hysterium</bold></italic>9. Phragmospores with swollen supra-median cell,
usually more than 3-septate, in one case with pigmented dictyospores and red
centrum pigmentation (<italic>Od.
pulchrum</italic>)...........................................................................................
<italic><bold>Oedohysterium</bold></italic></p></list-item><list-item><p>10. Dictyospores hyaline, +/- gelatinous sheath, or pigmented, but short,
less than 25 μm in length..................................
<italic><bold>Hysterobrevium</bold></italic>10. Dictyospores hyaline, +/- gelatinous
sheath, or pigmented, but longer than 25 μm, or very large didymospores
(<italic>O.
curtisii</italic>)...............................................................................................................................................................................................
11</p></list-item><list-item><p>11. Dictyospores, if hyaline, then longer than 25 μm, or if pigmented,
then measuring (22–)25–34(–45) x
(6–)8–12(–17) μm, with 7–11 transverse and
1–2 vertical septa, and no red pigment associated with the hamathecium
(<italic>Gp. subrugosa</italic>).......... <italic><bold>Gloniopsis</bold></italic>11.
Dictyospores pigmented, of different length, or if similar in length to
<italic>Gp. subrugosa</italic>, then tropical with red pigment associated with the
hamathecium, or very large didymospores (<italic>O.
curtisii</italic>)...........................................................................................
12</p></list-item><list-item><p>12. Dictyospores or large didymospores borne in conchate, mytilinidioid,
thin-walled, slerenchymatous, fragile
fruitbodies..................................................................................................................................................................
<italic><bold>Ostreichnion</bold>Note</italic>: The genus <italic>Ostreichnion</italic>,
previously in the <italic>Mytilinidiaceae</italic>, was transferred to the
<italic>Hysteriaceae</italic> (<xref ref-type="bibr" rid="ref18">Boehm <italic>et
al.</italic> 2009</xref>).
12. Dictyospores borne in thick-walled,
navicular
hysterothecia...................................................................................................................
<bold>13</bold></p></list-item><list-item><p>13. Dictyospores pigmented, borne in typical hysterothecia, that are
erumpent or sessile on the substrate.....................
<italic><bold>Hysterographium</bold>Note</italic>: The genus <italic>Hysterographium</italic>,
with the type species <italic>Hg. fraxini</italic>, has been transferred out of the
<italic>Hysteriaceae</italic> as <italic>Pleosporomycetidae gen. incertae sedis</italic>(<xref ref-type="bibr" rid="ref18">Boehm <italic>et al.</italic> 2009</xref>).
Residual species classified as <italic>Hysterographium</italic>, for which sequence
data are lacking, are provisionally retained within the genus.
13.
Hysterothecia borne within the substrate, hardly erumpent, with cristate
longitudinal apex instead of a sulcus;
neotropical..........................................................................................................................................................................
<italic><bold>Hysterocarina</bold></italic></p></list-item></list></p><p>The genus <italic>Encephalographa</italic> was originally placed in the
<italic>Hysteriaceae</italic> by Renobales & Aguirre
(<xref ref-type="bibr" rid="ref92">1990</xref>) who thought it to be
lichenicolous. Tretiach & Modenesi
(<xref ref-type="bibr" rid="ref113">1999</xref>) demonstrated it to be
lichenised, and maintained its placement within the <italic>Hysteriaceae</italic>. The
latter authors illustrate endolithic, saxicolous, dichotomously branched,
laterally anastomosed, lirelliform pseudothecia with a longitudinal sulcus,
and clavate bitunicate asci bearing pigmented didymospores, highly reminiscent
of the saxicolous forms of <italic>Glonium circumserpens</italic>, in the
<italic>Gloniaceae</italic>. We recently were able to obtain fresh material of
<italic>Encephalographa elisae</italic> from Mauro Tretiach (Dipartimento di Biologia,
Università di Trieste, Trieste, Italy), and, although cultures failed,
we were able to isolate DNA directly from the ascomata (EB 0347 / BPI 879773).
Sequence data presented here indicate that <italic>E. elisae</italic> does not reside
within the <italic>Hysteriaceae</italic>, nor within the <italic>Gloniaceae</italic>. Instead,
<italic>E. elisae</italic> lies outside of the <italic>Pleosporomycetidae</italic> and
<italic>Dothideomycetidae</italic> (<xref ref-type="fig" rid="fig1">Fig.
1</xref>).</p><p>To summarise, we accept the following genera in the <italic>Hysteriaceae</italic>:
<italic>Actidiographium, Gloniella, Gloniopsis, Hysterium, Hysterobrevium,
Hysterocarina, Oedohysterium, Ostreichnion, Psiloglonium</italic>, and
<italic>Rhytidhysteron</italic>. Dichotomous keys are presented here for hysteriaceous
fungi, with the caveat that phylogenetically unrelated taxa share the same
key. Thus, despite their transference from the <italic>Hysteriaceae</italic>(<xref ref-type="bibr" rid="ref18">Boehm <italic>et al.</italic> 2009</xref>),
the genera <italic>Hysterographium, Farlowiella, Glonium</italic> and
<italic>Anteaglonium</italic> (<xref ref-type="bibr" rid="ref81">Mugambi &
Huhndorf 2009</xref>), are nevertheless included in the key. This is
because they typically possess ascomata that have traditionally been referred
to as hysterothecia.</p><p><italic><bold>Hysterium</bold></italic> Tode, Schriften Berlin. Ges. Naturf. Freunde 5:
53 (1784).</p><p>The genus <italic>Hysterium</italic> is characterised by pigmented versicolorous or
concolorous asymmetric phragmospores, three- or more transversely-septate,
borne in hysterothecia. A historical overview of the nomenclature of the genus
was presented in Boehm <italic>et al.</italic>(<xref ref-type="bibr" rid="ref18">2009</xref>). Zogg
(<xref ref-type="bibr" rid="ref122">1962</xref>) recognised two
morphological types within the genus. Type I is characterised by 3-septate
phragmospores, and includes the versicolorous type species <italic>H.
pulicare</italic> (<xref ref-type="fig" rid="fig2">Fig.
2A–B</xref>), and its closely related concolorous counterpart,
<italic>H. angustatum</italic> (<xref ref-type="fig" rid="fig2">Fig.
2C–F</xref>), both extremely common in the temperate zones of both
hemispheres. These are followed by <italic>H. vermiforme</italic>(<xref ref-type="fig" rid="fig2">Fig. 2G–K</xref>), from Africa,
and the much larger-spored <italic>H. macrosporum</italic>, reported from North
America and China (<xref ref-type="bibr" rid="ref111">Teng
1933</xref>). Although Zogg
(<xref ref-type="bibr" rid="ref122">1962</xref>) did not accept <italic>H.
hyalinum</italic>, Lohman (<xref ref-type="bibr" rid="ref60">1934</xref>)
provided legitimacy to the epithet, noting that pigmentation is delayed in the
maturation of the 3-septate ascospores
(<xref ref-type="bibr" rid="ref18">Boehm <italic>et al.</italic>2009</xref>).</p><p>Type II corresponds to a different phragmospore, one in which, typically,
there are five or more septa, and in which there exists a swollen cell, either
just above the median septum (<italic>i.e.</italic>, supramedian) or, rarely, some
distance up from the median septum. Type II includes, by increasing spore
length, the cosmopolitan <italic>H. insidens</italic>(<xref ref-type="fig" rid="fig3">Fig. 3A–D</xref>), the
larger-spored counterpart <italic>H. sinense</italic>(<xref ref-type="fig" rid="fig3">Fig. 3E–H</xref>), and the
unusual <italic>H. magnisporum</italic>, 7-septate, with three of the septa crowded to
each end, the two central cells much larger. The latter two species are
reported from China (<xref ref-type="bibr" rid="ref111">Teng
1933</xref>) and North America (Boehm, unpubl. data). <italic>Hysterium
velloziae</italic>, provisionally included by Zogg
(<xref ref-type="bibr" rid="ref122">1962</xref>), with up to 21 septa
at maturity, has only been reported from Africa
(<xref ref-type="bibr" rid="ref51">van der Linde 1992</xref>).</p><p><fig position="float" id="fig2"><label>Fig. 2.</label><caption><p>The genus <italic>Hysterium</italic> (Clade C). A–B. <italic>Hysterium
pulicare</italic> [<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123377&link_type=cbs">CBS
123377</ext-link> (BPI 878723), U.S.A.]; C–F. <italic>Hysterium
angustatum</italic> [ANM 120 (ILLS), U.S.A.; not incl.]; G–K. <italic>Hysterium
vermiforme</italic> [GKM 1234 (BPI 879785), Kenya]; L–Q. <italic>Hysterium
barrianum</italic> <italic>sp. nov.</italic> [ANM 1495 (ILLS 59908 = holotype), U.S.A.].
Scale bar (habitat) = 500 μm; Scale bar (spores and asci) = 20 μm.</p></caption><graphic xlink:href="49fig2"></graphic></fig></p><p>An additional two species have been recently described. <italic>Hysterium
asymmetricum</italic> (<xref ref-type="bibr" rid="ref21">Checa <italic>et al.</italic>2007</xref>) from Costa Rica, has outer centrum tissues pigmented red,
and 3-septate phragmospores, showing an extended basal cell. <italic>Hysterium
andinense</italic> has been recently described from the conifer <italic>Austrocedrus
chilensis</italic> in Argentina (<xref ref-type="bibr" rid="ref75">Messuti &
Lorenzo 1997</xref>). However, molecular and morphological data
(<xref ref-type="bibr" rid="ref18">Boehm <italic>et al.</italic> 2009</xref>)
has placed this taxon in the <italic>Mytilinidiaceae</italic>, as <italic>Mytilinidion
andinense</italic>, based on <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123562&link_type=cbs">CBS
123562</ext-link> (BPI 878737). This brings the total number of species
within the genus <italic>Hysterium</italic> to 10. An additional new species is
described here.</p><p><italic><bold>Hysterium barrianum</bold></italic> E.W.A. Boehm, A.N. Miller, G.K.
Mugambi, S.M. Huhndorf & C.L. Schoch, <bold>sp. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515330&link_type=mb">MB515330</ext-link>.
<xref ref-type="fig" rid="fig2">Fig. 2L–Q</xref>.</p><p><italic>Etymology</italic>: Named after the late Dr Margaret E. Barr, preeminent
American mycologist.</p><p>Ascomata inconspicue hysterothecioidea, modice compressa e latere in parte
superiore, paulo conniventia, sulco inconspicuo angusto, latera paucis striis
profundis praedita; ascomata recta vel flexuosa, sessilia, raro furcata,
matura altiora quam lata, 1–2.5 mm longa, 250–450 μm alta,
200–300 μm lata. Pseudoparaphyses hyalinae, cellulares, 1–2
μm latae, supra ascos ramosae epithecium formantes. Asci bitunicati,
cylindrici, breviter stipitati, (110–)125–135 x 15–20 μm.
Phragmosporae fusiformes, angustae, rectae vel paulo curvatae, primum
hyalinae, maturae pallide luteae, quaque cellula guttulis magnis
refringentibus repleta, (7–)9(–11)-septatae,
(35–)40–45(–55) x (7–)9–10(–12) μm.</p><p><italic>Ascomata</italic> atypically hysterithecioid, somewhat laterally compressed
in the upper region, slightly connivent, sulcus very shallow, existing as a
narrow rim, sides laterally striate, striae few and deep, straight to
flexuous, sessile on the substrate, rarely bifurcating, taller than wide at
maturity: 1–2.5 mm long x 250–450 μm high, 200–300 μm
wide. <italic>Pseudoparaphyses</italic> hyaline, cellular, 1–2 μm wide,
branched above the ascal layer to form an epithecium. <italic>Asci</italic>bitunicate, cylindrical, short-stipitate, (110–)125–135 x
15–20 μm (n = 9). <italic>Phragmospores</italic> fusiform, narrow, hyaline
and straight when young, becoming pale-yellow to lightly clear-brown, and
curved when mature, highly guttulate, with guttulae large, highly refractive,
present in every cell, with (7–)9(–11) septa, measuring
(35–)40–45(–55) x (7–)9–10(–12) μm when
mature (n = 27).</p><p><italic>Specimens examined</italic>: <bold>U.S.A.</bold>, Tennessee, Sevier Co., Great
Smoky Mountains National Park, Elkmont, Little River Trail, 35° 39' 13.4''
N, 83° 34' 44.7'' W, 686 m elev., 5 Nov. 2007, A.N. Miller, S.M. Huhndorf,
J.L. Crane, T.J. Atkinson, I. Promputtha, M. Grief, G.K. Mugambi, & P.
Chaudhary, deposited as ILLS 59908 (ANM 1495) = <bold>holotype</bold>; BPI 879783 =
<bold>paratype</bold>; Tennessee, Sevier Co., Great Smoky Mountains National Park,
Chimney Tops Picnic Area, Cove Hardwood Loop Trail, 35° 38' 10.7'' N,
83° 29' 32.1'' W, 4 Nov. 2007, A.N. Miller, S.M. Huhndorf, J.L. Crane,
T.J. Atkinson, I. Promputtha, M. Grief, G.K. Mugambi & P. Chaudhary,
deposited as ILLS 59907 (ANM 1442), and BPI 879784.</p><p><italic>Notes</italic>: A superficial resemblance exists between <italic>Hysterium
barrianum</italic> in Clade C, with <italic>H. sinense</italic> in Clade D. The
phragmospores of <italic>H. barrianum</italic>(<xref ref-type="fig" rid="fig2">Fig. 2N–Q</xref>) have a
similar number of septa, (7–)9(–11), as those of <italic>H.
sinense</italic> (<xref ref-type="fig" rid="fig3">Fig. 3H</xref>), the
latter with (3–)5–9(–11) septa. The two species also have
spores of similar length. However, the width measurements of <italic>H.
barrianum</italic>, (35–)40–45(–55) x
(7–)9–10(–12) μm, serve to separate it from <italic>H.
sinense</italic>, (34–)38–50 x 11–15 μm. Most importantly,
<italic>H. barrianum</italic> does not possess a swollen or tumid supra-median cell,
as does <italic>H. sinense</italic> and the closely related <italic>H. insidens</italic>.
Furthermore, <italic>H. barrianum</italic> is highly guttulate, and lightly pigmented
at maturity, whereas <italic>H. sinense</italic> and <italic>H. insidens</italic> possess few
if any guttulae, and are much darker in pigmentation at maturity. Lastly,
molecular data place the species in different groups within the
<italic>Hysteriaceae</italic>.</p><p>In this study, we were able to secure a wide taxon sampling strategy for
the genus <italic>Hysterium</italic> (<xref ref-type="table" rid="tbl1">Table
1</xref>), including multiple isolates for seven of the eleven currently
recognised species, namely: <italic>H. pulicare</italic> (1), <italic>H. angustatum</italic>(7), <italic>H. vermiforme</italic> (1), <italic>H. insidens</italic> (2), <italic>H. sinense</italic>(2), <italic>H. barrianum</italic> (2) and <italic>H. hyalinum</italic> (1). Multiple gene
phylogenies indicate that the genus <italic>Hysterium</italic> is polyphyletic, along
three separate lines, two within the <italic>Hysteriaceae</italic> and one, <italic>H.
hyalinum</italic>, outside of the family (<xref ref-type="fig" rid="fig1">Fig.
1</xref>). This implies that the evolution of pigmented phragmospores
borne in hysterothecia has occurred at least three times within the
<italic>Pleosporomycetidae</italic>.</p><p>Sequence data indicate that Clade C contains the type species,
<italic>Hysterium pulicare</italic>, as well as the closely related <italic>H.
angustatum</italic>, and <italic>H. vermiforme</italic>(<xref ref-type="fig" rid="fig1">Fig. 1</xref>). All three taxa have
3-septate, pigmented phragmospores, corresponding to Type I. Also, within
Clade C resides the newly described <italic>H. barrianum</italic>, with 9-sepate
spores. None of these species has a swollen supra-median cell. Accessions of
<italic>H. angustatum</italic>, originating from South Africa (CMW 20409), Kenya (GKM
243A), New Zealand (SMH 5211.0, SMH 5216) and the United States, New Jersey
(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123334&link_type=cbs">CBS 123334</ext-link>) and
Wisconsin (<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=236.34&link_type=cbs">CBS
236.34</ext-link>), form a highly supported monophyletic clade with <italic>H.
pulicare</italic>, collected from the United States, New York
(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123377&link_type=cbs">CBS 123377</ext-link>). Both
species possess similar pigmented 3-septate phragmospores, versicolorous in
<italic>H. pulicare</italic> and concolorous in <italic>H. angustatum</italic>. Interestingly,
∼10 % of the ascospores within a given hysterothecium of <italic>H.
pulicare</italic> are typically found to be concolorous
(<xref ref-type="bibr" rid="ref16">Bisby 1941</xref>). Likewise,
versicolorous ascospores have also been observed in <italic>H. angustatum</italic>,
stated at less than ∼5 % for a given hysterothecium
(<xref ref-type="bibr" rid="ref49">Lee & Crous 2003</xref>).
Although ascospore size in <italic>H. pulicare</italic> may be twice that found in
<italic>H. angustatum</italic> (<xref ref-type="bibr" rid="ref122">Zogg
1962</xref>), a certain degree of overlap in spore length measurements
exists between the two, and molecular data presented here and elsewhere
(<xref ref-type="bibr" rid="ref18">Boehm <italic>et al.</italic> 2009</xref>)
indicate that they are closely related.</p><p><fig position="float" id="fig3"><label>Fig. 3.</label><caption><p>The genus <italic>Oedohysterium</italic> (Clade D). A–D. <italic>Oedohysterium
insidens</italic> [ANM 1443 (BPI 879799), U.S.A.]; E–H. <italic>Oedohysterium
sinense</italic> [ANM 119 (ILLS), U.S.A.; not incl.]. Scale bar (habitat) = 500
μm; Scale bar (spores and asci) = 20 μm.</p></caption><graphic xlink:href="49fig3"></graphic></fig></p><p>In this study, one of the <italic>H. angustatum</italic> accessions from Tennessee
(ANM 85), did not cluster with the other surveyed <italic>H. angustatum</italic> in
Clade C. Instead, ANM 85 clustered with <italic>H. vermiforme</italic> from Kenya (GKM
1234). Spore measurements of ANM 85 (ILLS) were compared to the other <italic>H.
angustatum</italic> accessions from the United States
(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123334&link_type=cbs">CBS 123334</ext-link> / BPI
878724), Kenya (GKM 243A, EA), and New Zealand (SMH 5211.0, F) which formed
the other sub-clade within Clade C. All of these specimens showed remarkably
little variability in their spore morphology. Additionally, no obvious
differences were noted in their fruitbody morphology. This may indicate early
stages of speciation within the taxon, with sequence variation preceding
morphologic change.</p><p>Grouping with the anomalous <italic>H. angustatum</italic> ANM 85, was <italic>H.
vermiforme</italic>, a taxon known only from the original description by Massee in
1901 from West Africa (Ghana). The isolate included here (GKM 1234 / BPI
879785; Fig. G–K) originated from Mt. Kenya, Kenya, and possesses
smaller spore measurements, (20–)25–28 x (4–)5–6
μm, than those given by Massee
(<xref ref-type="bibr" rid="ref74">1901</xref>), and reiterated by Zogg
(<xref ref-type="bibr" rid="ref122">1962</xref>), as
(30–)35–40 x 12–14 μm. In other respects, however, BPI
879785 matches closely Massee's
(<xref ref-type="bibr" rid="ref74">1901</xref>) original description,
and we choose here to simply expand the spore measurements for <italic>H.
vermiforme</italic> to (20–)25–40 x (4–)5–14 μm, rather
than describe a new species.</p><p>The 3-septate <italic>H. hyalinum</italic>(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=237.34&link_type=cbs">CBS 237.34</ext-link>) lies
outside of the <italic>Hysteriaceae</italic> altogether. It falls in a small,
isolated, but well-supported clade along with the type species of
<italic>Hysterographium</italic>, namely <italic>Hg. fraxini</italic>. Since only one isolate
is represented, it is premature to draw conclusions. Molecular data indicate
that the remaining two species of <italic>Hysterium</italic> in our survey, namely
<italic>H. sinense</italic> and <italic>H. insidens</italic>, are not related to the type
<italic>H. pulicare</italic> and associated species within Clade C. Rather, data
indicate that they belong to Clade D. As such, we propose the following new
genus to accommodate these taxa.</p><p><italic><bold>Oedohysterium</bold></italic> E.W.A. Boehm & C.L. Schoch, <bold>gen.
nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515421&link_type=mb">MB515421</ext-link>.</p><p><italic>Etymology</italic>: Greek, <italic>Oedo-</italic> meaning swollen, referring to the
swollen supra-median cell of the ascospores and <italic>Hys</italic>- from
<italic>Hysterium.</italic></p><p>Hysterothecia solitaria vel gregaria, iuvenia erumpentia, deinde
superficialia, navicularia, nonnumquam linearia, plus minusve parallela, neque
confluentia, nonnumquam angulo inserta, raro flexuosa vel furcata, plerumque
utrinque obtuse, et fissura longitudinali prominente praedita. Latitudo
altitudine minor vel major. Peridium crassum, carbonaceum, maturum fragile,
per longitudinem striatum, basim versus incrassatum, sursum attenuatum,
bistratosum. Pseudoparaphyses cellulares, 1–2.5 μm latae, hyalinae,
septatae, sursum ramosae, vulgo epithecium pigmentatum ascos obtegens
formantes. Asci cylindrici vel clavati, bitunicati. Ascosporae irregulariter
biseriatae, phragmoseptatae (dictyoseptatae), fusiformes, curvatae, utrinque
angustatae, ad septum medium constrictae, (4–)6–8(–11)
septis divisae, primum pallide luteae, deinde brunnescentes. Cellula (raro duo
cellulae) ascosporarum supramediana conspicue inflata. Anamorphe ad
<italic>Septonema</italic> pertinens.</p><p><italic>Hysterothecia</italic> isolated to gregarious, erumpent when young,
superficial when mature, navicular, sometimes linear in more or less parallel
rows, but non confluent laterally, or sometimes situated at angles, rarely
flexuous or bifurcating, usually with obtuse ends, and with a prominent
longitudinal slit. Sometimes, taller than wide, other times wider than tall.
<italic>Peridium</italic> thick, carbonaceous, brittle with age, longitudinally
striated on the margins, thickened towards base, less thick apically, composed
of two to three distinct layers, the inner compressed and pallid, the outer
thickened and pigmented. <italic>Pseudoparaphyses</italic> cellular, 1–2.5 μm
wide, hyaline, septate, branched above, forming a usually pigmented epithecium
above the asci. <italic>Asci</italic> cylindrical to clavate, usually short stipitate,
and bitunicate. <italic>Ascospores</italic> irregularly biseriate in ascus, typically
phragmospores, in one case dictyospores, curved, fusiform, with tapering
apices, constricted at the median septum, with (4–)6–8(–11)
septa, at first hyaline-yellow, then pigmented sepia to brown at maturity.
Genus characterised by a swollen or tumid supra-median cell, rarely with two
cells swollen. <italic>Anamorph</italic>: <italic>Septonema</italic>.</p><p><italic>Type species</italic>: <italic>Oedohysterium insidens</italic> (Schwein.) E.W.A.
Boehm & C.L. Schoch, comb. nov.</p><p><italic><bold>Oedohysterium insidens</bold></italic> (Schwein.) E.W.A. Boehm & C.L.
Schoch, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515422&link_type=mb">MB515422</ext-link>.
<xref ref-type="fig" rid="fig3">Fig.
3A–D</xref>.
<italic>Basionym</italic>: <italic>Hysterium insidens</italic>Schwein., Trans. Amer. Philos. Soc., New Series 4(2): 244. 1832.</p><p><list list-type="simple"><list-item><p>≡ <italic>Hysterographium insidens</italic> (Schwein.) Sacc., Syll. Fung. 2:
778. 1883.</p></list-item><list-item><p>= <italic>Hysterium complanatum</italic> Duby, Mém. Soc. Phys. Genève
16(1): 38. 1862.</p></list-item><list-item><p>= <italic>Hysterium depressum</italic> Berk. & M.A. Curtis, Grevillea 4(29):
10. 1875.</p></list-item><list-item><p>= <italic>Hysterium fusigerum</italic> Berk. & M.A. Curtis, Grevillea 4(29):
11. 1875 (as `<italic>fusiger'</italic>).</p></list-item><list-item><p>= <italic>Hysterium berengeri</italic> Sacc., Syll. Fung. 2: 751. 1883.</p></list-item><list-item><p>= <italic>Hysterium janusiae</italic> Rehm, Hedwigia 37: 299. 1898.</p></list-item><list-item><p>= <italic>Hysterium apiculatum</italic> Starbäck, Bidrag Kungl. Svenska
Vetensk.-Akad. Hist. 25(1): 19. 1899.</p></list-item><list-item><p>= <italic>Hysterium batucense</italic> Speg., Revista Fac. Agron. Univ. Nac. La
Plata 6(1): 116. 1910.</p></list-item><list-item><p>= <italic>Hysterium andicola</italic> Speg., Anal. Mus. Nac. Hist. Nat. B. Aires
23: 85. 1912.</p></list-item><list-item><p>= <italic>Hysterium atlantis</italic> Maire, Mém. Soc. Sci. Nat. Maroc. 45:
35. 1937.</p></list-item><list-item><p>= <italic>Hysterium lavandulae</italic> Urries, Ann. Jard. Bot. Madrid 1: 64.
1941.</p></list-item></list></p><p><italic>Hysterothecia</italic> isolated to gregarious, variably erumpent to
sessile, 0.5–2.5 mm long, 0.2–0.5 mm high, lying parallel, but not
confluent laterally, generally in line with the grain of the wood, and
striated laterally with age. <italic>Pseudoparaphyses</italic> hyaline, cellular,
1–2.0 μm wide, walls thickened at apices, forming an epithecium,
borne in mucilage, above the ascal layer, often encrusted with dark, pigmented
crystals. <italic>Asci</italic> bitunicate, cylindrical, 8-spored, irregularly
biseriate, 130–150 x 15–24 μm, short stipitate, and with a
prominent apical nasse, especially when young. <italic>Ascospores</italic>phragmospores transversely (4–)6–8(–11)-septate, constricted
at the median septum, inequilateral, slightly curved, at first hyaline-yellow,
then brown at maturity, with a prominent swollen supra-median cell. If
5-septate, then swollen cell located at the second position; if 6-septate,
then often the third from the top, measuring (20–)23–28(–38)
x (5–)7–10(–13) μm. Principally North- and South-America,
and Europe (Italy), from bark and old wood of <italic>Pinus, Larix, Castanea,
Quercus, Eucalyptus, Fraxinus, Aspidosperma,</italic> and <italic>Lavandula</italic>(<xref ref-type="bibr" rid="ref122">Zogg 1962</xref>). Also reported
from South Africa (<xref ref-type="bibr" rid="ref51">van der Linde,
1992</xref>). Anamorph: <italic>Septonema spilomeum</italic>.</p><p><italic><bold>Oedohysterium sinense</bold></italic> (Teng) E.W.A. Boehm & C.L.
Schoch, <bold>comb. nov</bold><italic>.</italic> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515423&link_type=mb">MB515423</ext-link>.
<xref ref-type="fig" rid="fig3">Fig.
3E–H</xref>.
<italic>Basionym</italic>: <italic>Hysterium sinense</italic> Teng,
Sinensia 4: 134. 1933.</p><p><list list-type="simple"><list-item><p>= <italic>Hysterium macrosporum</italic> Teng, Sinensia 4: 134. 1933, non Peck,
Rep. (Annual) New York State Mus. Nat. Hist. 26: 83. 1874 (1873).</p></list-item></list></p><p><italic>Hysterothecia</italic> scattered to subgregarious, linear, sometimes
parallel but non-confluent laterally, more often lying at irregular angles,
depending on the grain of the substrate, striated in age, usually of a similar
size (2–3.5 mm in length), that is, maturing synchronously in a given
colony. <italic>Pseudoparaphyses</italic> hyaline to pale-yellow, cellular,
2–2.5 μm wide, apically branched, walls of even thickness along
length, forming a darkened gelatinous epithecium above the ascal layer, +/-
encrusted with pigmented crystals. <italic>Asci</italic> bitunicate, cylindrical,
8-spored, irregularly biseriate, 140–170 x 26–30 μm,
short-stipitate, ascospores biseriate to subseriate in ascus, with a prominent
apical nasse, especially when young, but sometimes persisting through
maturity. <italic>Ascospores</italic> large, fusiform, asymmetric, curved
phragmospores, at first hyaline, then pale-yellow to -brown, finally deep
brown at maturity, with (3–)5–9(–11) septa, with a medial
septal constriction, measuring (34–)38–50 x 11–15 μm,
and, like <italic>Od. insidens</italic>, with a prominent swollen or tumid
supra-median cell, usually located just above the median septum. From North
America (Boehm, unpubl. data), Europe
(<xref ref-type="bibr" rid="ref122">Zogg 1962</xref>), China
(<xref ref-type="bibr" rid="ref111">Teng 1933</xref>), and South Africa
(<xref ref-type="bibr" rid="ref51">van der Linde 1992</xref>), on
decorticated hardwood trees and structures (<italic>e.g.</italic>, aged fence
posts).</p><p><italic>Notes</italic>: Species of <italic>Oedohysterium</italic> belonging to Clade D are
characterised by elongate asymmetric spores with more than 3 septa, typically
showing a swollen or tumid supra-median cell. In this study, two
single-ascospore isolates of <italic>Od. sinense</italic>, one from South Africa
(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123345&link_type=cbs">CBS 123345</ext-link> / BPI
878730), and one from the United States, New Jersey (EB 0339 / BPI 879800),
cluster with two isolates of <italic>Od. insidens</italic>, both from the United
States, Massachusetts (<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=238.34&link_type=cbs">CBS
238.34</ext-link>) and Tennessee (ANM 1443 / BPI 879799). Both species have
remarkably similar phragmospores (<italic>e.g.</italic>,
<xref ref-type="fig" rid="fig3">Fig. 3D</xref> <italic>versus</italic><xref ref-type="fig" rid="fig3">Fig. 3H</xref>). As these two taxa
belong to Clade D and are far removed from the type species, <italic>H.
pulicare</italic>, in Clade C, we propose that they be accommodated in the new
genus <italic>Oedohysterium</italic>. An additional new combination is proposed
below.</p><p><italic><bold>Oedohysterium pulchrum</bold></italic> (Checa, Shoemaker &
Umaña) E.W.A. Boehm & C.L. Schoch, <bold>comb. nov</bold><italic>.</italic>MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515424&link_type=mb">MB515424</ext-link>.</p><p><italic>Basionym</italic>: <italic>Hysterographium pulchrum</italic> Checa, Shoemaker &
Umaña, <italic>Mycologia</italic> 99: 289. 2007.</p><p><italic>Notes</italic>: The newly described <italic>Hg. pulchrum</italic> from Costa Rica
(<xref ref-type="bibr" rid="ref21">Checa <italic>et al.</italic> 2007</xref>)
also falls within Clade D (<xref ref-type="fig" rid="fig1">Fig.
1</xref>) and is here transferred to <italic>Oedohysterium</italic>, as <italic>Od.
pulchrum</italic> (DQ 402184 / DAOM 234345). This is because molecular data
indicate a close association with the two species of <italic>Oedohysterium, Od.
insidens</italic> and <italic>Od. sinense</italic>. At first surprising, on further
consideration, this sub-clade forms a natural assemblage premised on
morphological features. The spores of all three taxa show a remarkable degree
of similarity in morphology, which includes being similarly pigmented,
slightly curved and fusiform, with a common number of transverse septa. The
sole difference is the presence of one or two vertical septa in <italic>Od.
pulchrum</italic>, a feature noted by the authors to be absent in some spores
(<xref ref-type="bibr" rid="ref21">Checa <italic>et al.</italic> 2007</xref>).
Most importantly, like <italic>Od. insidens</italic> and <italic>Od. sinense, Od.
pulchrum</italic> also possesses a swollen supra-median cell. Interestingly, a
striking resemblance to the phragmospores of <italic>Od. insidens</italic> can be seen
for those spores of <italic>Od. pulchrum</italic> that do not possess vertical septa
(<xref ref-type="bibr" rid="ref21">Checa <italic>et al.</italic> 2007</xref>).
This is based on similarities in shape (<italic>e.g.</italic>, curved and fusiform),
size [(20–)23–28(–38) x (5–)7–10(–13)
μm <italic>versus</italic> 22–25(–27) x 5–6 μm], and in the
number of transverse septa (4–)6–8(–11) <italic>versus</italic>(5–)6, for <italic>Od. insidens</italic> and <italic>Od. pulchrum</italic>,
respectively. As molecular data indicate that the presence or absence of
vertical septa should be considered a sympleisiomorphic character state within
the <italic>Hysteriaceae</italic> (<xref ref-type="bibr" rid="ref18">Boehm <italic>et
al.</italic> 2009</xref>), we feel justified in including both phragmospores
and dictyospores within the genus <italic>Oedohysterium</italic>.</p></sec><sec><title>Key to the species of <italic>Hysterium</italic> and
<italic>Oedohysterium</italic></title><p><list list-type="simple"><list-item><p>1. Phragmospores mainly
3-septate.............................................................................................................................................................
2
1. Phragmospores concolorous, more than 3-septate, in one instance
pigmented dictyospores with 1-2 vertical septa (<italic>Od.
pulchrum</italic>)..........................................................................................................................................................................................
7</p></list-item><list-item><p>2. Phragmospores either versicolorous or delayed
concolorous..................................................................................................................
3
2. Phragmospores truly concolorous (sepia to dark brown in
colour)...........................................................................................................
4</p></list-item><list-item><p>3. Terminal cell mainly remaining hyaline with inner spore cells pigmented
brown (versicolorous); ascospores 20–40 x 6–12 μm;
cosmopolitan...........................................................................................................................
<italic><bold>H. pulicare</bold></italic>3. Phragmospores tardily pigmented, often
remaining hyaline for quite some time after discharge, but eventually becoming
uniformly concolorous; 20–26(–28) x 6–8.5 μm; North
America.................................................... <italic><bold>H.
hyalinum</bold>Note</italic>: Currently recognised as <italic>Pleosporomycetidae sp.
incertae sedis</italic> (<xref ref-type="bibr" rid="ref18">Boehm <italic>et al.</italic>2009</xref>).</p></list-item><list-item><p>4. Phragmospores 3-septate, 28 μm or less in
length..................................................................................................................................
5
4. Phragmospores 3-septate, longer than 28
μm..........................................................................................................................................
6</p></list-item><list-item><p>5. Phragmospores (12–)14–21(–28) x
(3–)4–8(–10) μm, firmly 3-septate, no septal
constrictions; end-cells obtuse;
cosmopolitan......................................................................................................................................................................
<italic><bold>H. angustatum</bold></italic>5. Phragmospores
(14–)15–18(–20) x 5–7 μm; 3- (rarely 2- or
4)-septate; prominently constricted at first-formed septum; basal cell
extended; red hamathecial pigment;
neotropical..........................................................................................
<italic><bold>H. asymmetricum</bold></italic></p></list-item><list-item><p>6. Phragmospores fusoid, slightly curved, guttulate; (20–)25–40
x (4–)5–14 μm; West and East
Africa............................... <italic><bold>H. vermiforme</bold></italic>6.
Phragmospores fusoid, curved, highly guttulate; 40–57 x 11–15
μm; on <italic>Pinus</italic>, North America and China......... <italic><bold>H.
macrosporum</bold></italic> <bold>Peck</bold></p></list-item><list-item><p>7. Phragmospores or dictyospores (4-) 6- to 8- (11-) celled, fusiform in
outline, with +/- swollen supra-median cell(s)...............................
8
7. Phragmospores with more than 11 septa, fusiform, pale brown,
(13–)14–15(–21)-septate, (35–)45–50(–60) x
(10–)12–13(–14) μm;
Africa......................................................................................................................
<italic><bold>H. velloziae</bold></italic></p></list-item><list-item><p>8. Swollen supra-median cell(s) present, either phragmospores or
dictyospores
(<italic>Oedohysterium</italic>).............................................................
9
8. Phragmospores only, no swollen supra-median cells(s)
present............................................................................................................
11</p></list-item><list-item><p>9. Dictyospores lightly pigmented, 22–25(–27) x 5–6
μm, with (5–)6 transverse and 1 vertical septum in either cell or
both cells adjacent to the primary septum, absent in some spores, with a
swollen supra-median cell; typically with red pigment in the hamathecium;
neotropical (Costa
Rica)...........................................................................
<italic><bold>Od. pulchrum</bold></italic>9. With no red pigment
present...................................................................................................................................................................
10</p></list-item><list-item><p>10. Phragmospores with (4–)6–8(–11) septa, slightly
curved, fusiform, at first hyaline-yellow then reddish brown at maturity, if
5-septate, showing a swollen cell at the second position, if 6-septate, often
the third from the top, +/- median septal constriction,
(20–)23–28(–38) x (5–)7–10(–13) μm;
cosmopolitan...........................................................
<italic><bold>Od. insidens</bold></italic>10. Phragmospores larger, fusiform, straight
to curved, at first hyaline, then yellow or pale brown, finally deep brown;
swollen supra-median cell(s) present, (3–)5–9(–11) septa,
with median septal constriction; (34–)38–50 x 11–15 μm;
cosmopolitan..................................................................................................................................
<italic><bold>Od. sinense</bold></italic></p></list-item><list-item><p>11. Phragmospores fusiform, narrow, straight to very slightly curved, pale
hyaline at first, then pale-yellow at maturity, with highly refractive
guttules, in every cell, with (7–)9(–11) septa, no supra-median
swollen cell(s), (35–)40–45(–55) x
(7–)9–10(–12) μm; North
America..........................................................................................................
<italic><bold>H. barrianum</bold></italic>11. Phragmospores oblong, wide, slightly
curved, bulging on one side, nearly hyaline and 1-septate at first, becoming
clear brown and 7-septate, septa highly asymmetric, (2–)3 of the septa
close to each end, the two central cells much larger; 48–67 x
15–20 μm; China and North
America......................................................... <italic><bold>H.
magnisporum</bold></italic></p></list-item></list></p><p>We choose to provide the following dichotomous key whereby all
hysteriaceous fungi, bearing transversely septate pigmented phragmospores
(including <italic>Od. pulchrum</italic> with dictyospores) are identified together,
with the caveat that unrelated taxa appear in the same key.</p><p><italic><bold>Gloniella</bold></italic> Sacc., Syll. Fung. 2: 765 (1883).</p><p>The genus <italic>Gloniella</italic> was established by Saccardo
(<xref ref-type="bibr" rid="ref96">1883</xref>) to accommodate
hysteriaceous fungi that possess hyaline phragmospores, from 3- to 9-septate.
Zogg (<xref ref-type="bibr" rid="ref122">1962</xref>) recognised six
species: three collected on ferns from Europe and the Mediterranean, namely
<italic>Gl. adianti</italic> on <italic>Adiantum</italic>, and <italic>Gl. graphidioidea</italic> and
<italic>Gl. normandina</italic>, both on <italic>Pteridium</italic>. Zogg also accepted
<italic>Gl. sardoa</italic> from <italic>Populus</italic> in Europe, <italic>Gl. typhae</italic> on
<italic>Typha</italic>, the latter described from Europe
(<xref ref-type="bibr" rid="ref122">Zogg 1962</xref>) and Chile
(<xref ref-type="bibr" rid="ref62">Lorenzo & Messuti 1998</xref>),
and <italic>Gl. bambusae</italic> on <italic>Bambusa</italic> from Brazil. Since then, an
additional three species have been described: <italic>Gl. gracilis</italic> from Costa
Rica (<xref ref-type="bibr" rid="ref21">Checa <italic>et al.</italic>2007</xref>), <italic>Gl. corticola</italic> from India
(<xref ref-type="bibr" rid="ref85">Pande & Rao 1991</xref>), and
<italic>Gl. clavatispora</italic> from South Africa
(<xref ref-type="bibr" rid="ref109">Steinke & Hyde 1997</xref>).
More recently, Barr (<xref ref-type="bibr" rid="ref10">2009</xref>)
recognised <italic>Gl. abietina</italic> on <italic>Abies</italic> from Idaho, and <italic>Gl.
lapponica</italic> on <italic>Arctostaphylos</italic> from Washington. A number of species
in the key may be conspecific, since reported spore measurements are identical
or nearly so.</p></sec><sec><title>Key to the species of <italic>Gloniella</italic></title><p><list list-type="simple"><list-item><p>1. Ascospores 3-septate, shorter than 15
μm...............................................................................................................................................
2
1. Ascospores 3- or more-septate, and
longer..............................................................................................................................................
3</p></list-item><list-item><p>2. Ascospores 10–15 x 5–6 μm;
India........................................................................................................................................
<italic><bold>Gl. corticola</bold></italic>2. Ascospores 12–14 x 4–5 μm;
on <italic>Typha</italic>,
Europe......................................................................................................................
<italic><bold>Gl. typhae</bold></italic></p></list-item><list-item><p>3. On ferns in
Europe....................................................................................................................................................................................
4
3. Not on
ferns..............................................................................................................................................................................................
6</p></list-item><list-item><p>4. Ascospores (2–)3(–4)-septate,
(11–)15–20(–23) x 3–5 μm; on <italic>Adiantum</italic>,
Europe..................................................................
<italic><bold>Gl. adianti</bold></italic>4. Ascospores (3–)5(–7)-septate,
slightly
longer..........................................................................................................................................
5</p></list-item><list-item><p>5. Ascospores (3–)5-septate, (15–)18–20(–22) x
4–5 μm; on <italic>Pteridium</italic>,
Europe............................................................. <italic><bold>Gl.
graphidoidea</bold></italic>5. Ascospores 5–7-septate,
(22–)25–27(–30) x 3–4 μm; on <italic>Pteridium</italic>,
Europe..................................................................
<italic><bold>Gl. normandina</bold></italic></p></list-item><list-item><p>6. Ascospores 1–3-septate, 36–39 x 10 μm; on
<italic>Arctostaphylos</italic>, Western North
America....................................................... <italic><bold>Gl.
lapponica</bold></italic>6. Ascospores with more
septa.....................................................................................................................................................................
7</p></list-item><list-item><p>7. Ascospores 3(–5) septate, 20–27 um x 7–8 μm; on
<italic>Abies grandis</italic>, Western North
America................................................. <italic><bold>Gl.
abietina</bold></italic>7. Ascospores with more
septa.....................................................................................................................................................................
8</p></list-item><list-item><p>8. Ascospores (6–)7(–8)-septate,
(16–)18–21(–26) x 6–7(–8) μm; on
<italic>Populus</italic>,
Europe..............................................................
<italic><bold>Gl. sardoa</bold></italic>8. Ascospores
larger.....................................................................................................................................................................................
9</p></list-item><list-item><p>9. Ascospores (5–)6(–8)-septate, (18–)37(–41) x
10–11.5 μm, hyaline, smooth; on <italic>Avicennia marina</italic>, South
Africa........ <italic><bold>Gl. clavatispora</bold></italic>9. Ascospores smaller,
neotropical.............................................................................................................................................................
10</p></list-item><list-item><p>10. Ascospores 6–7-septate, 32–37(–40) x 4–6 μm;
Costa
Rica..................................................................................................
<italic><bold>Gl. gracilis</bold></italic>10. Ascospores (5–)6–7-septate,
(28–)32–38(–44) x (3–)4–8(–9) μm; on
<italic>Bambusa</italic>, Brazil....................................................
<italic><bold>Gl. bambusae</bold></italic></p></list-item></list></p><p><italic><bold>Hysterographium</bold></italic> Corda, Icon. Fung. 5: 34. 1842.</p><p><list list-type="simple"><list-item><p>= <italic>Hysteriopsis</italic> Speg., Revista Fac. Agron. Univ. Nac. La Plata 2:
308. 1907.</p></list-item><list-item><p>= <italic>Polhysterium</italic> Speg., Anales Mus. Nac. Buenos Aires 23: 87.
1912.</p></list-item><list-item><p>= <italic>Fragosoa</italic> Cif., in Ciferri & Fragoso, Bol. Real Soc. Esp.
Hist. Nat., Secc. Biol. 26(3-4): 194. 1926.</p></list-item></list></p><p>Although the genus <italic>Hysterographium</italic> has been removed from the
<italic>Hysteriaceae</italic> (<xref ref-type="bibr" rid="ref18">Boehm <italic>et
al.</italic> 2009</xref>), and is currently recognised as
<italic>Pleosporomycetidae gen. incertae sedis</italic>, it is included here. This is
because it forms the basis for a number of new combinations within the family.
The genus is characterised by pigmented dictyospores, with one to several
longitudinal septa, ovoid to ellipsoid-fusoid, relatively broad, usually
constricted at the first-formed septum. Zogg
(<xref ref-type="bibr" rid="ref122">1962</xref>) extensively revised
the synonymy of the genus and accepted four species: <italic>Hysterographium
flexuosum</italic> (<xref ref-type="fig" rid="fig4">Fig.
4A–B</xref>) and <italic>Hg. fraxini</italic>(<xref ref-type="fig" rid="fig4">Fig. 4C–D</xref>), the type,
both with large dictyospores, prominently constricted at the median septum,
the former with slightly longer, narrower spores. Zogg
(<xref ref-type="bibr" rid="ref122">1962</xref>) also accepted <italic>Hg.
mori</italic> and <italic>Hg. subrugosum</italic>, with smaller, fewer-celled
dictyospores, short and squat in the former, longer and more slender in the
latter, both also constricted at the median septum.</p><p>Since then, an additional three species have been described:
<italic>Hysterographium minus</italic> from Japan
(<xref ref-type="bibr" rid="ref1">Amano 1983</xref>), <italic>Hg.
spinicola</italic> from South Africa, recollected from the thorns of
<italic>Acacia</italic> and validated by van der Linde
(<xref ref-type="bibr" rid="ref51">1992</xref>), with a brick-red
epithecium and spores only slightly longer than those of <italic>Hg. mori</italic>,
and, lastly, <italic>Hg. pulchrum</italic> from Costa Rica, also with a red pigment in
the hamathecium (<xref ref-type="bibr" rid="ref21">Checa <italic>et al.</italic>2007</xref>), here transferred to <italic>Oedohysterium</italic>, as <italic>Od.
pulchrum</italic>.</p><p><fig position="float" id="fig4"><label>Fig. 4.</label><caption><p>The genus <italic>Hysterographium</italic>. A–B. <italic>Hysterographium
flexuosum</italic> (EB 0098, U.S.A.; not incl.); C–D. <italic>Hysterographium
fraxini</italic> (EB 0100, U.S.A.; not incl.). Scale bar (habitat) = 1 mm; Scale
bar (spores) = 20 μm.</p></caption><graphic xlink:href="49fig4"></graphic></fig></p><p>Four of the seven species were surveyed in the present study, with multiple
isolates (<xref ref-type="table" rid="tbl1">Table 1</xref>):
<italic>Hysterographium mori</italic> (8), <italic>Hg. subrugosum</italic> (3), <italic>Hg.
fraxini</italic> (2) and <italic>Od. pulchrum</italic> (1), falling into no fewer than
three separate clades, two within the <italic>Hysteriaceae</italic> (Clades A and D)
and one far removed from the family (<xref ref-type="fig" rid="fig1">Fig.
1</xref>). The latter clade includes the type species for the genus
<italic>Hysterographium</italic>, namely <italic>Hg. fraxini</italic>, represented by isolates
from Switzerland (<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=109.43&link_type=cbs">CBS
109.43</ext-link>), deposited by Zogg in 1943, and from Canada
(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=242.34&link_type=cbs">CBS 242.34</ext-link>),
deposited by Lohman in 1934. <italic>Hysterographium fraxini</italic> forms a
well-supported clade distant from the <italic>Hysteriaceae,</italic> but remains
within the <italic>Pleosporomycetidae</italic>(<xref ref-type="fig" rid="fig1">Fig. 1</xref>). As this is
substantiated by two geographically disparate isolates from two different
continents, deposited by two reputable workers, it is significant. The
implication is that the genus <italic>Hysterographium</italic> must follow the type
species and be removed from the <italic>Hysteriaceae</italic>(<xref ref-type="bibr" rid="ref18">Boehm <italic>et al.</italic> 2009</xref>).
Species with pigmented dictyospores remaining within the
<italic>Hysteriaceae</italic>, previously classified in <italic>Hysterographium</italic>, must
therefore be accommodated in other genera. In this study, these would include
the following species, for which we have sequence data: <italic>Hysterographium
mori, Hg. subrugosum</italic>, and <italic>Hg. pulchrum</italic> (= <italic>Od.
pulchrum</italic>). The remaining species for which we do not have sequence data,
namely <italic>Hg. minus, Hg. spinicola</italic> and <italic>Hg. flexuosum</italic>, must
remain as species of <italic>Hysterographium</italic>, until such time that sequence
data are available. We therefore propose the following new genus.</p><p><italic><bold>Hysterobrevium</bold></italic> E.W.A. Boehm & C.L. Schoch, <bold>gen.
nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515329&link_type=mb">MB515329</ext-link>.</p><p><italic>Etymology</italic>: <italic>Hystero-</italic> from <italic>Hysterographium</italic>, Latin
<italic>brevis</italic>, short, referring to the spores of the type, <italic>Hb.
mori</italic>.</p><p>Hysterothecia navicularia, fissura longitudinali prominente praedita,
utrinque acuminata vel obtusa, linearia vel flexuosa, solitaria vel gregaria,
vulgo per longitudinem striata, nonnumquam erecta, quasi stipitata,
superficialia vel partim in substrato immersa. Asci bitunicati, cyindrici vel
clavati. Dictyosporae pigmentatae vel hyalinae, plerumque breviores quam 25
μm, ad septum medium constrictae; ascosporae hyalinae vel luteae iuvenes
vulgo strato mucido circumdatae; pigmentatae pallide brunneae, pariete levi;
ascosporae ovoideae vel obovoideae, apice obtuso vel acuminato,
3–4(–6) septis transversalibus et 1–2 longitudinalibus
divisae.</p><p><italic>Hysterothecia</italic> navicular, with a prominent longitudinal slit,
variable with acuminate to obtuse ends, linear to flexuous, solitary to
densely gregarious, surface usually longitudinally striate, sometimes erect,
superficial, almost stipitate, to erumpent and partially embedded in
substrate, the latter especially when gregarious. <italic>Asci</italic> bitunicate,
cylindrical to clavate. <italic>Ascospores</italic> pigmented or hyaline dictyospores,
usually less than 25 μm long, constricted at least at the median septum. If
hyaline to pale-yellow, then typically associated with a gelatinous sheath
when young, dissipating with age. If pigmented then lightly so, transparent
clear brown, walls smooth; ascospores generally ovoid to obovoid, with either
obtuse or acuminate ends, 3–4(–6) transverse septa, and 1–2
longitudinal septa, these mostly associated with the two central cells, but
highly variable and sometimes at oblique angles in the end cells.</p><p><italic>Type species</italic>: <italic>Hysterobrevium mori</italic> (Schwein.) E.W.A. Boehm
& C.L. Schoch, comb. nov.</p><p><italic><bold>Hysterobrevium mori</bold></italic> (Schwein.) E.W.A. Boehm & C.L.
Schoch, <bold>comb. nov</bold><italic>.</italic> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515335&link_type=mb">MB515335</ext-link>.
<xref ref-type="fig" rid="fig5">Fig.
5J–R</xref>.
<italic>Basionym</italic>: <italic>Hysterium mori</italic>Schwein., Trans. Amer. Philos. Soc. 4(2): 244. 1832.</p><p><list list-type="simple"><list-item><p>≡ <italic>Hysterographium mori</italic> (Schwein.) Rehm, Ascomyceten No. 363.
1876.</p></list-item><list-item><p>= <italic>Hysterium grammodes</italic> De Not., Giorn. Bot. Ital. 2 (7–8):
55. 1847.</p></list-item><list-item><p>≡ <italic>Hysterographium grammodes</italic> (De Not.) Sacc., Syll. Fung. 2:
782. 1883.</p></list-item><list-item><p>= <italic>Hysterium rousselii</italic> De Not., Piren. Ister. 2(7–8): 19.
1847.</p></list-item><list-item><p>≡ <italic>Hysterographium rousselii</italic> (De Not.) Sacc., Syll. Fung. 2:
779. 1883.</p></list-item><list-item><p>= <italic>Hysterium vulgare</italic> De Not., Piren. Ister. 2(7–8): 18.
1847.</p></list-item><list-item><p>= <italic>Hysterium australe</italic> Duby, Mém. Soc. Phys. Genève
16(1): 44. 1862.</p></list-item><list-item><p>= <italic>Hysterium lesquereuxii</italic> Duby, Mém. Soc. Phys.
Genève 16(1): 41. 1862.</p></list-item><list-item><p>≡ <italic>Hysterographium lesquereuxii</italic> (Duby) Sacc., Syll. Fung. 2:
779. 1883.</p></list-item><list-item><p>= <italic>Hysterium gerardi</italic> Cooke & Peck, Bull. Buffalo Soc. Nat. Sci.
3: 33. 1875.</p></list-item><list-item><p>≡ <italic>Hysterographium gerardi</italic> (Cooke & Peck) Sacc., Syll.
Fung. 2: 783. 1883.</p></list-item><list-item><p>= <italic>Hysterium viticolum</italic> Cooke & Peck, Bull. Buffalo Soc. Nat.
Sci. 3: 33. 1875.</p></list-item><list-item><p>≡ <italic>Hysterographium viticola</italic> (Cooke & Peck) Rehm, Ascomyc.
No. 316, in Sacc., Syll. Fung. 2: 782. 1883.</p></list-item><list-item><p>= <italic>Hysterium variabile</italic> Cooke & Peck, Bull. Buffalo Soc. Nat.
Sci. 3: 33. 1875.</p></list-item><list-item><p>≡ <italic>Hysterographium variabile</italic> (Cooke & Peck) Sacc., Syll.
Fung. 2: 780. 1883.</p></list-item><list-item><p>= <italic>Hysterium formosum</italic> Cooke, in Harkness & Cooke, Grevillea 7:
3. 1878.</p></list-item><list-item><p>≡ <italic>Hysterographium formosum</italic> (Cooke) Sacc., Syll. Fung. 2:
783. 1883.</p></list-item><list-item><p>= <italic>Hysterium putaminum</italic> Cooke, Grevillea 7: 48. 1878.</p></list-item><list-item><p>≡ <italic>Hysterographium putaminum</italic> (Cooke) Sacc., Syll. Fung. 2:
783. 1883.</p></list-item><list-item><p>= <italic>Hysterographium portenum</italic> Speg., Anales Soc. Ci. Argent., Secc.
Santa Fe. 9(4): 185. 1880.</p></list-item><list-item><p>= <italic>Hysterographium grammodes</italic> var. <italic>minus</italic> Sacc., Syll. Fung.
2: 783. 1883.</p></list-item><list-item><p>= <italic>Hysterographium pumilionis</italic> Rehm, Discom. 1(3): 21. 1887.</p></list-item><list-item><p>= <italic>Hysterographium guaraniticum</italic> Speg., Anales Soc. Ci. Argent.,
Secc. Santa Fe. 26(1): 56. 1888.</p></list-item><list-item><p>= <italic>Hysterographium punctiforme</italic> Pat., Bull. Soc. Mycol. France 4:
120. 1888.</p></list-item><list-item><p>= <italic>Hysterographium ruborum</italic> Cooke, in Rehm, Ascom., No. 918.
1888.</p></list-item><list-item><p>= <italic>Hysterium insulare</italic> P. Karst. & Har., Rev. Mycol. Toulouse
No. 47: 1890.</p></list-item><list-item><p>= <italic>Hysterographium incisum</italic> Ellis & Everh., Bull. Torrey Bot.
Club 24: 462. 1897.</p></list-item><list-item><p>= <italic>Hysterographium ziziphi</italic> Pat., Cat. Rais. Pl. Cell. Tunisie: 112.
1897 (as “<italic>zizyphi</italic>”).</p></list-item><list-item><p>= <italic>Hysterographium rousselii</italic> (De Not.) Sacc. var. <italic>piri</italic>Feltgen, Vorst. Pilz. Luxemb. Nachtr. 3: 111. 1903.</p></list-item></list></p><p><italic>Hysterothecia</italic> erumpent-superficial, ellipsoidal, oblong, linear or
cylindrical, 1–2(–3.5) mm long, 220–275(–440) μm
wide, by 190–330 μm high, mostly straight and lying parallel, but not
confluent laterally, often gregarious and crowded so as to cover the
substrate, longitudinally striate in age, navicular with tapering ends. Two
types of hysterothecial aggregations regularly observed, depending on
substrate: (1) Colonies on weathered, whitened decorticated hardwood often
forming large oval colonies, with acuminate ends, measuring 5–15 cm in
length, with hysterothecia gregarious in the center, densely packed in
longitudinal formations, showing multiple stages of development, and darkening
the adjacent substrate; when young, prior to emergence of hysterothecia,
smaller colonies are seen, but still presenting darkened oval patches, often
with coelomycetous anamorph present. (2) Colonies on bark (<italic>i.e.</italic>,
corticolous) less gregarious, not darkening the substrate, hysterothecia often
situated at angles, rather than in parallel orientation. <italic>Peridium</italic>30–60 μm thick medially, to 100+ μm at the base, distinctly
three-layered in cross-section, the outer layer darkly pigmented, the middle
less so, and the inner layer, thin-walled, pallid and compressed.
<italic>Pseudoparaphyses</italic> cellular, septate, persistent, 1–2 μm wide,
hyaline, thickened apically, branched and forming an epithecium in a
gelatinous matrix above the ascal layer. <italic>Asci</italic> cylindrical to clavate,
bitunicate, short-stipitate, (50–)80–110 x 10–18 μm.
<italic>Ascospores</italic> pigmented, thin-walled dictyospores, obovoid, ends obtuse,
3–(5–7)-septate, with 1–2(–3) vertical septa usually
associated with mid-cells, but on occasion also present obliquely in end
cells, constricted at the median septum, sometimes, when fully hydrated, at
additional, more distal septa, measuring (12–)14–22(–26) x
(5–)7–10(–11) μm. <italic>Anamorph</italic> coelomycetous,
<italic>Aposphaeria</italic>-like in nature, in culture conidiomata as irregular
locules, with conidiogenous cells 8–10 x 1.5–2 μm;
<italic>conidia</italic> (2–)2.5–3.5(–4) x 1–2 μm (Lohman
1932). Cosmopolitan, on aged, usually decorticated, weathered wood or bark of
<italic>Pinus, Juniperus, Salix, Ostrya, Castanea, Quercus, Ulmus, Morus, Pyrus,
Amelanchier, Crataegus, Rubus, Cercocarpus, Prunus, Gleditsia,</italic> various
<italic>Fabaceae, Melia, Pistacia, Cotinus, Rhus, Acer, Ziziphus, Vitis, Fraxinus,
Olea,</italic> and <italic>Aspidosperma</italic>(<xref ref-type="bibr" rid="ref122">Zogg 1962</xref>).</p><p><italic><bold>Hysterobrevium smilacis</bold></italic> (Schwein.) E.W.A. Boehm & C.L.
Schoch, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515336&link_type=mb">MB515336</ext-link>.
<xref ref-type="fig" rid="fig5">Fig.
5F–I</xref>.
<italic>Basionym</italic>: <italic>Hysterium smilacis</italic>Schwein., Schriften Naturf. Ges. Leipzig 1: 49. 1822.</p><p><list list-type="simple"><list-item><p>≡ <italic>Gloniopsis smilacis</italic> (Schwein.) Underw. & Earle, Bull.
Alabama Agric. Exp. Sta. 80: 196. 1897.</p></list-item><list-item><p>≡ <italic>Hysterographium smilacis</italic> (Schwein.) Ellis & Everh., N.
Amer. Pyrenomyc. 709. 1892.</p></list-item><list-item><p>= <italic>Hysterium biforme</italic> Fr., Observ. Mycol. (Havniae) 2: 354.
1818.</p></list-item><list-item><p>≡ <italic>Gloniopsis biformis</italic> (Fr.) Sacc., Syll. Fung. 2: 773.
1883.</p></list-item><list-item><p>= <italic>Hysterium elongatum β curvatum</italic> Fr., Elench. Fung.
(Greifswald) 2: 138. 1828.</p></list-item><list-item><p>= <italic>Hysterium curvatum</italic> Fr., Elench. Fung. 2: 139. 1828.</p></list-item><list-item><p>≡ <italic>Gloniopsis curvata</italic> (Fr.) Sacc., Syll. Fung. 2: 775.
1883.</p></list-item><list-item><p>= <italic>Hysterium rocheanum</italic> Duby, Mém. Soc. Phys. Genève
16: 51. 1862.</p></list-item><list-item><p>≡ <italic>Gloniopsis rocheana</italic> (Duby) Sacc., Syll. Fung. 2: 773.
1883.</p></list-item><list-item><p>= <italic>Hysterographium naviculare</italic> P. Karst., Symb. Mycol. Fenn. 6: 37.
1877.</p></list-item><list-item><p>= <italic>Hysterium gloniopsis</italic> W.R. Gerard in Peck, Rep. New York State
Mus. 32: 49. 1877 (1879).</p></list-item><list-item><p>≡ <italic>Hysterographium gloniopsis</italic> (W.R. Gerard) Ellis &
Everh., N. Amer. Pyrenomyc. 708. 1892.</p></list-item><list-item><p>≡ <italic>Gloniopsis gloniopsis</italic> (W.R. Gerard) House, Bull. New York
State Mus. 219-220: 235. 1920.</p></list-item><list-item><p>= <italic>Gloniella scortechiniana</italic> Sacc. & Roum., Rev. Mycol. Toulouse
5: tab. 41, fig. 17. 1883.</p></list-item><list-item><p>= <italic>Gloniopsis gerardiana</italic> Sacc., Syll. Fung. 2: 774. 1883.</p></list-item><list-item><p>= <italic>Gloniopsis decipiens</italic> var. <italic>cisti</italic> Rehm, Hedwigia 25: 13.
1886.</p></list-item><list-item><p>= <italic>Gloniopsis cisti</italic> Rehm, Hedwigia 25: 13. 1896.</p></list-item><list-item><p>= <italic>Gloniopsis ambigua</italic> Sacc., Ann. Mycol. 10(3): 317. 1912.</p></list-item><list-item><p>= <italic>Gloniopsis ellisii</italic> Cash, Mycologia 31: 294. 1939.</p></list-item></list></p><p><fig position="float" id="fig5"><label>Fig. 5.</label><caption><p>The genus <italic>Hysterobrevium</italic> (Clade A). A–E. <italic>Hysterobrevium
constrictum</italic> [SMH 5211.1 (F), New Zealand]; F–I. <italic>Hysterobrevium
smilacis</italic> [GKM 426N (EA), Kenya]; L–N. <italic>Hysterobrevium mori</italic>[SMH 5273 (BPI 879787), U.S.A.]; O–R. <italic>Hysterobrevium mori</italic> [ANM
43 (ILLS), U.S.A.; not incl]. Scale bar (habitat) = 500 μm; Scale bar
(spores and asci) = 10 μm.</p></caption><graphic xlink:href="49fig5"></graphic></fig></p><p><italic>Hysterothecia</italic> erumpent, many times surrounded at the base by
ruptured epidermis or periderm, especially when borne in herbaceous stems,
much less so on wood, then completely superficial, 0.5–1.5 mm long,
300–400 μm wide, 200–250 μm high, longitudinally striated.
<italic>Peridium</italic> 25–50 μm wide, narrower at base within the
substrate, widest at mid-point, carbonaceous and brittle when dry.
<italic>Pseudoparaphyses</italic> cellular, septate, persistent, 1–1.5 μm
wide, hyaline to pale yellow in mass, branched above, forming an epithecium,
but not darkly pigmented, exposed surface yellow-brown. <italic>Asci</italic>cylindrical to clavate, bitunicate, short-stipitate, 70–120 x
15–25 μm at maturity. <italic>Ascospores</italic> asymmetric, hyaline to pale
yellow dictyospores, with acuminate ends, and a gelatinous sheath that usually
dissipates at maturity, measuring (13–)15–26(– 31) x
(4–)5–9(–10) μm. Spore septation highly variable, usually
3–5(–9)-septate and with 1(–3) vertical septa, passing
through multiple mid-cells, and usually prominently constricted at the median
septum, when fresh and hydrated, sometimes constriced along multiple
transverse septa. <italic>Anamorph</italic> coelomycetous, <italic>Aposphaeria</italic>-like.
Cosmopolitan on <italic>Pinus, Chamaerops, Smilax, Populus, Salix, Juglans,
Betula, Fagus, Quercus, Ficus, Pyrus, Crataegus, Rubus, Rosa, Prunus, Robinia,
Butea, Pistacia, Cotinus, Acer, Cistus, Erica,</italic> and <italic>Lavandula</italic>(<xref ref-type="bibr" rid="ref122">Zogg 1962</xref>).</p><p><italic>Notes</italic>: <italic>Hysterobrevium mori</italic>, while falling within the
<italic>Hysteriaceae</italic>, finds itself in two separate clades
(<xref ref-type="fig" rid="fig1">Fig. 1</xref>). In Clade A, one set
of North American <italic>Hb. mori</italic> isolates associates with six highly
geographically diverse isolates of <italic>Hb. smilacis</italic>. The <italic>Hb.
mori</italic> isolates originate from the United States, from New Jersey
(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123336&link_type=cbs">CBS 123336</ext-link>,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123564&link_type=cbs">CBS 123564</ext-link>), New
York (<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123335&link_type=cbs">CBS 123335</ext-link>,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123563&link_type=cbs">CBS 123563</ext-link>),
Indiana (SMH 5273) and Michigan (SMH 5286). The <italic>Hb. smilacis</italic> isolates
originate from the United States, from Indiana (SMH 5280) and Michigan
(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=200.34&link_type=cbs">CBS 200.34</ext-link>), as
well as from South Africa (CMW 18053), Sweden
(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=114601&link_type=cbs">CBS 114601</ext-link>) and
Kenya (GKM 426N). Dictyospores of both species are of similar shape, size and
degree of septation: (12–)14–22(–26) x
(5–)7–10(–11) μm, 3–(5–7)-septate, with
1–2(–3) vertical septa, for <italic>Hb. mori versus</italic> (13–)
15–26(–31) x (4–)5–9(–10) μm,
3–5(–9)-septate, with 1(–3) vertical septa, for <italic>Hb.
smilacis</italic>. They differ in the absence of pigmentation and the presence of
a gelatinous sheath in the latter. Thus, these two species, previously
classified in two separate genera, <italic>Hysterographium</italic> and
<italic>Gloniopsis</italic>, are in fact closely related, with each species far
removed from the type species of their respective genera. Further support for
this argument, can be found in Lohman
(<xref ref-type="bibr" rid="ref58">1933a</xref>), who found a similar
<italic>Aposphaeria</italic> anamorph for both <italic>Hb. mori</italic> (as <italic>Hg.
mori</italic>) and <italic>Hb. smilacis</italic> (as <italic>Gp. gerardiana</italic>) and stated
that they were indistinguishable in culture. The implication is that both taxa
should be united within the same genus, for which we propose
<italic>Hysterobrevium</italic>.</p><p>In addition to the association with <italic>Hb. smilacis</italic> in Clade A,
<italic>Hb. mori</italic> also finds itself in Clade D. As this is validated by two
geographically diverse isolates, one from the United States, Michigan
(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=245.34&link_type=cbs">CBS 245.34</ext-link>) and
one from Kenya (GKM 1013 / BPI 879788), it is significant. Spore measurements
of the Kenyan accession GKM 1013 (BPI 879788) in Clade D <italic>versus</italic> those
of other <italic>Hb. mori</italic> accessions in Clade A, represented by SMH 5273 /
BPI 879787, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123335&link_type=cbs">CBS
123335</ext-link> / BPI 878734, and
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123336&link_type=cbs">CBS 123336</ext-link> / BPI
878733, failed to detect any significant morphological differences; nor were
there any appreciable differences detected in their hysterothecia. The
association of <italic>Hb. mori</italic> with unrelated taxa within the
<italic>Hysteriaceae</italic> in Clade A and D may be significant in that <italic>Hb.
mori</italic> has long been regarded as a highly variable taxon
(<xref ref-type="bibr" rid="ref32">Ellis & Everhart 1892</xref>,
<xref ref-type="bibr" rid="ref58">Lohman 1933a</xref>), resulting in
the synonymy of no fewer than 30 names since its inception by Schweinitz in
1832 (<xref ref-type="bibr" rid="ref122">Zogg 1962</xref>). Future
studies may well reveal that <italic>Hb. mori</italic> contains a number of cryptic
species, morphologically similar, but genetically unrelated. We propose an
additional new combination below.</p><p><italic><bold>Hysterobrevium constrictum</bold></italic> (N. Amano) E.W.A. Boehm &
C.L. Schoch, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515337&link_type=mb">MB515337</ext-link>.
<xref ref-type="fig" rid="fig5">Fig. 5A–E</xref>.</p><p><italic>Basionym</italic>: <italic>Gloniopsis constricta</italic> N. Amano, Trans. Mycol.
Soc. Japan 24: 289. 1983.</p><p><italic>Notes</italic>: Amano (<xref ref-type="bibr" rid="ref1">1983</xref>)
described a small-spored species of <italic>Gloniopsis</italic> from Japan, <italic>Gp.
constricta</italic>, noting a prominent median septal constriction. The
measurements of the dictyospores were given as 10.4–13.2 x 4.4–5.8
μm, usually with 3–4 transverse and one vertical septum that passes
through one to three cells. Although not mentioned
(<xref ref-type="bibr" rid="ref1">Amano 1983</xref>), the illustrations
depict a very thick wall and dictyospores highly symmetric in outline and
septation. Amano (<xref ref-type="bibr" rid="ref1">1983</xref>) stated
of the spores “...hyaline, later becoming brown...”, but did not
mention the presence of a gelatinous sheath. He also noted that the closest
resemblance is with <italic>Hb. smilacis</italic> (as <italic>Gp. curvata</italic>), the
latter however with slightly larger spores. In this study, we were fortunate
to obtain a specimen from New Zealand (SMH 5211.1, deposited in F;
<xref ref-type="fig" rid="fig5">Fig. 5A–E</xref>) that
corresponds to the published description given by Amano
(<xref ref-type="bibr" rid="ref1">1983</xref>), but differs on several
counts. Like <italic>Gp. constricta</italic>, the hyaline dictyospores in SMH 5211.1,
are highly symmetric and thick-walled, (1–)3(–4)-septate, with
1(–2) vertical septa, but the constriction at the median septum in SMH
5211.1, while present, is not prominent. Also unlike <italic>Gp. constricta</italic>,
the spores in SMH 5211.1 have an obvious gelatinous sheath when young, but
this quickly dissipates with age, and may be completely absent in mature
specimens. In SMH 5211.1, the spores measure (18–)20(–23) x
10–12 μm, which is considerably larger than those of <italic>Gp.
constricta</italic>. Nevertheless, these differences, in our opinion, are not
sufficient to warrant a new species, and we choose here to simply expand the
spore measurements to (11–)13–20(–23) x 5–12 μm,
rather than describe a new species, proposing instead the new combination
<italic>Hb. constrictum</italic>.</p><p><italic><bold>Gloniopsis</bold></italic> De Not., Giorn. Bot. Ital. 2(2): 23. 1847.</p><p>A review of the nomenclatural history of the genus <italic>Gloniopsis</italic> was
given in Boehm <italic>et al.</italic>(<xref ref-type="bibr" rid="ref18">2009</xref>). The genus is
characterised by hyaline to yellow dictyospores, often inequilateral, curved,
in outline obovoid, ends obtuse to sub- to acuminate, multi-septate, with one
or more longitudinal septa, constricted at the first-formed septum, sometimes
constricted at additional septa, and usually surrounded by a gelatinous
sheath, which may dissipate with age. Zogg
(<xref ref-type="bibr" rid="ref122">1962</xref>) synonymised a number
of names under the type species, <italic>Gp. praelonga</italic>(<xref ref-type="fig" rid="fig6">Fig. 6A–B</xref>), and accepted
only one additional species, namely <italic>Gp. curvata</italic> with smaller
ascospores. Barr (<xref ref-type="bibr" rid="ref8">1990a</xref>)
proposed to include this latter species under the earlier name <italic>Gp.
smilacis</italic>, following Cash
(<xref ref-type="bibr" rid="ref20">1939</xref>). In this study, we have
transferred <italic>Gp. smilacis</italic> to <italic>Hysterobrevium</italic>, closely related
to <italic>Hb. mori</italic> in Clade A. Recently, <italic>Gp. argentinensis</italic>,
previously considered by Zogg
(<xref ref-type="bibr" rid="ref122">1962</xref>) as a doubtful species,
was reinstated by Lorenzo & Messuti
(<xref ref-type="bibr" rid="ref62">1998</xref>). The authors state that
the ascospores are 7-septate, with 1–3(–4) longitudinal septa,
some passing through multiple cells, in outline widely ellipsoid, measuring
20–26 x 9–12 μm. The septation and spore measurements are
nearly identical to those of <italic>Gp. praelonga</italic>, the latter
5–7(–10)-septate, with 2–3 longitudinal septa,
(16–)20–32(–34) x (6–)9–12(–15) μm. We
therefore synonymise <italic>Gp. argentinensis</italic> under <italic>Gp. praelonga</italic>.
Lastly, Amano (<xref ref-type="bibr" rid="ref1">1983</xref>) described
an additional two species of <italic>Gloniopsis</italic> from Japan, namely <italic>Gp.
macrospora</italic> and <italic>Gp. constricta</italic>, the latter transferred here to
<italic>Hysterobrevium</italic> (Clade A).</p><p>Molecular data indicate that the genus <italic>Gloniopsis</italic> is polyphyletic,
with the type, <italic>Gp. praelonga</italic>, belonging to Clade D
(<xref ref-type="fig" rid="fig1">Fig. 1</xref>). Closely associated
with the type, are a number of species possessing pigmented dictyospores,
which would previously have been classified in the genus
<italic>Hysterographium</italic> (<italic>e.g., Hysterographium subrugosum</italic>). Based on
molecular data presented here, we therefore propose to emend the genus
<italic>Gloniopsis</italic>, to include both hyaline and pigmented dictyospores. The
following new combination is proposed, as well as two new species from
Africa.</p><p><fig position="float" id="fig6"><label>Fig. 6.</label><caption><p>The genus <italic>Gloniopsis</italic> (Clade D). A–B. <italic>Gloniopsis
praelonga</italic> [<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123337&link_type=cbs">CBS
123337</ext-link> (BPI 878725), U.S.A.]; C–F. <italic>Gloniopsis
subrugosa</italic> [GKM 1214 (BPI 879776), Kenya]; G. <italic>Gloniopsis
subrugosa</italic> (<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123346&link_type=cbs">CBS
123346</ext-link>, BPI 878735; South Africa). Scale bar (habitat) = 500
μm; Scale bar (spores and asci) = 20 μm.</p></caption><graphic xlink:href="49fig6"></graphic></fig></p><p><italic><bold>Gloniopsis subrugosa</bold></italic> (Cooke & Ellis) E.W.A. Boehm &
C.L. Schoch, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515338&link_type=mb">MB515338</ext-link>.
<xref ref-type="fig" rid="fig6">Fig.
6C–G</xref>.
<italic>Basionym</italic>: <italic>Hysterium subrugosum</italic>Cooke & Ellis, Grevillea 5: 54. 1876.</p><p><list list-type="simple"><list-item><p>≡ <italic>Hysterographium subrugosum</italic> (Cooke & Ellis) Sacc.,
Syll. Fung. 2: 780. 1883.</p></list-item><list-item><p>= <italic>Hysterographium hiascens</italic> Rehm, Ber. Naturhist. Vereins. Augsburg
26: 780. 1881.</p></list-item><list-item><p>= <italic>Hysterographium kansense</italic> Ellis & Everh., Erythea 2: 22.
1894.</p></list-item><list-item><p>= <italic>Hysterographium cylindrosporum</italic> Rehm, Bih. Kongl. Svenska
Vetensk.-Akad. Handl. 25(6): 11. 1899.</p></list-item><list-item><p>= <italic>Hysterographium minutum</italic> M.L. Lohman, Pap. Michigan Acad. Sci.
17: 267. 1933.</p></list-item></list></p><p><italic>Hysterothecia</italic> erumpent to superficial, scattered to densely
crowded, navicular, straight to flexuous, with tapered ends, surface not
striated in age, but smooth to sub-rugose in texture, 1–2 mm long,
250–350 μm diam. <italic>Peridium</italic> composed of small
pseudoparenchymatous cells, heavily pigmented at the surface, not showing a
distinct number of layers, relatively smooth on outer surface.
<italic>Pseudoparaphyses</italic> narrowly cellular, septate, 1–1.5 μm in
diam., hyaline, branched above the asci, borne in a gelatinous matrix.
<italic>Asci</italic> cylindrical to clavate, bitunicate, short-stipitate,
80–150 x 18–25 μm, with a prominent apical nasse, especially
when young. <italic>Ascospores</italic> pigmented thin-walled, dictyospores
(22–)25–34(–45) x (6–)8–12(–17) μm,
mostly with 7–11 transverse and 1–2 vertical septa, hardly
constricted at septa, clear brown, ends paler at times, slightly asymmetric in
outline. <italic>Anamorph</italic> coelomycetous, <italic>Aposphaeria-</italic>like
(<xref ref-type="bibr" rid="ref58">Lohman 1933a</xref>). Less
frequently collected, but reported from North America
(<xref ref-type="bibr" rid="ref9">Barr 1990b</xref>), Europe
(<xref ref-type="bibr" rid="ref122">Zogg 1962</xref>), Argentina
(<xref ref-type="bibr" rid="ref76">Messuti & Lorenzo 2003</xref>)
and from South Africa (<xref ref-type="bibr" rid="ref51">van der Linde
1992</xref>) as well. Old wood and bark of <italic>Populus, Quercus, Celtis,
Crataegus, Rosa,</italic> and <italic>Cotinus</italic>(<xref ref-type="bibr" rid="ref122">Zogg 1962</xref>), as well as on
weathered fence posts and old planks (Boehm, unpubl. data).</p><p><italic>Notes</italic>: In the current study, we were able to include three
geographically diverse isolates of <italic>Gp. praelonga</italic>(<xref ref-type="table" rid="tbl1">Table 1</xref>), two from South
Africa (<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=112415&link_type=cbs">CBS 112415</ext-link>and CMW 19983 / PREM 57539), and one from the United States, New Jersey
(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123337&link_type=cbs">CBS 123337</ext-link> / BPI
878725). These isolates cluster together in Clade D and associate with one
isolate of <italic>Gp. subrugosa</italic> from South Africa
(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123346&link_type=cbs">CBS 123346</ext-link> / BPI
878735). Both <italic>Gp. praelonga</italic> and <italic>Gp. subrugosa</italic> are somewhat
similar in the shape, size and septation of their dictyospores, hyaline in the
former (<xref ref-type="fig" rid="fig6">Fig. 6B</xref>), pigmented in
the latter (<xref ref-type="fig" rid="fig6">Fig. 6G</xref>). The
spores of <italic>Gp. praelonga</italic> are (16–)20–32(–34) x
(6–)9–12(–15) μm, and those of <italic>Gp. subrugosa</italic> are
(22–)25–34(–45) x (6–)8–12(–17) μm.
Septation is also similar in both species, with 5–7(–10)
transverse and 2–3 vertical septa in <italic>Gp. praelonga</italic> and
7–11 transverse and 1–2 vertical septa in <italic>Gp. subrugosa</italic>.
They differ in pigmentation and the presence of a gelatinous sheath in the
type. Molecular data indicate that they are closely related.</p><p>An additional two isolates of <italic>Gp. subrugosa</italic>, from Kenya (GKM 1214
/ BPI 879776) and Cuba (SMH 557 / BPI 879777), are more distantly related and
do not fall in Clade D. Moreover, no morphological differences were noted
between these two more distantly associated isolates of <italic>Gp. subrugosa</italic>and <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123346&link_type=cbs">CBS 123346</ext-link> (BPI
878735) from South Africa in Clade D. Although spore morphology dictates that
all three specimens of <italic>Gp. subrugosa</italic> should be classified as the same
species, molecular data point to genetic heterogeneity within the taxon. This
is similar to the situation in <italic>Hb. mori</italic>, mentioned earlier, which,
despite identical morphologies, finds affinities in both Clades A and D.
<italic>Hysterobrevium mori</italic> and, to a lesser extent, <italic>Gp. subrugosa</italic>,
may represent ancestral lineages that have maintained stable morphologies,
while simultaneously incurring sufficient genetic change to, in the case of
<italic>Hb. mori</italic>, fall into different clades within the family.
Alternatively, these isolates may represent examples of convergent evolution
among genetically unrelated lineages, which produce remarkably similar
ascospores and hysterothecia. Also associating with <italic>Gp. praelonga</italic> and
<italic>Gp. subrugosa</italic> in Clade D are two new species from East Africa,
described below.</p><p><italic><bold>Gloniopsis arciformis</bold></italic> E.W.A. Boehm, G.K. Mugambi, S.M.
Huhndorf & C.L. Schoch, <bold>sp. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515331&link_type=mb">MB515331</ext-link>.
<xref ref-type="fig" rid="fig7">Fig. 7A–H</xref>.</p><p><italic>Etymology</italic>: Latin <italic>arcus</italic>, a bow or arch, referring to the
arcuate or arciform dictyospores.</p><p>Hysterothecia solitaria vel pauca aggregata, recta vel flexuosa,
carbonacea, plerumque erecta, conspicue applanata et altiora quam lata,
(0.5–)1–2.5 mm longa, 250–350 μm lata, 400–600
μm alta, per longitudinem striata, sulco inconspicuo maturitate clauso.
Peridium 40–75 μm crassum in medio, basim versus crassius, sursum
tenuius, bistratosum. Pseudoparaphyses cellulares 1–1.5 μm latae,
ramosae, sursum magis crassitunicatae, epithecium pigmentatum ascos obtegens
formantes. Asci cylindrici vel clavati, stipite sinuoso, bitunicati,
50–75 × 14–18 μm; ascosporae irregulariter biseriatae,
dictyosporae, pigmentatae, tenuitunicatae, fragiles, facile dilabentes,
conspicue arcuatae, 3–5(–7)-septatae, 1–2(–3) septis
verticalibus divisae; cellulis centralibus multo maioribus quam distales, ad
septa haud constrictae, (10–)12–18(–22) x 6–10
μm.</p><p><italic>Hysterothecia</italic> solitary to sparsely aggregated, straight to
flexuous, carbonaceous, mainly erect, distinctly flattened and taller than
wide, (0.5–)1–2.5 mm long, 250–350 μm wide, by
400–600 μm high, longitudinally striated, with an inconspicuous
sulcus remaining closed at maturity. <italic>Peridium</italic> 40–75 μm thick
medially, thicker towards the base, thinner towards the sulcus, composed of
two layers, the inner thin, compressed and hyaline, the outer denser, and
darkly pigmented. <italic>Pseudoparaphyses</italic> cellular 1–1.5 μm wide,
branched and thicker-walled distally towards the top, forming a pigmented
epithecium above the asci. <italic>Asci</italic> cylindrical to clavate, with a
sinuous stalk, bitunicate, 50–75 x 14–18 μm (n = 7), ascospores
irregularly biseriate. <italic>Ascospores</italic> pigmented, thin-walled,
dictyospores, fragile, easily breaking under the slightest pressure,
pronouncedly arcuate or bent (arciform), and thus highly asymmetric,
3–5(–7)-septate, with 1–2(–3) vertical septa, these
mostly associated with the mid cells, which are much larger and swollen than
the end-cells, no septal constrictions, measuring (10–)
12–18(–22) x 6–10 μm (n = 17).</p><p><fig position="float" id="fig7"><label>Fig. 7.</label><caption><p>The genus <italic>Gloniopsis</italic> (Clade D). A–H. <italic>Gloniopsis
arciformis</italic> <italic>sp. nov.</italic> [GKM L166A (BPI 879774 = holotype), Kenya];
I–M. <italic>Gloniopsis kenyensis</italic> <italic>sp. nov.</italic> [GKM 1010 (BPI
879775 = holotype), Kenya]. Scale bar (habitat) = 500 μm; Scale bar (spores
and asci) = 10 μm.</p></caption><graphic xlink:href="49fig7"></graphic></fig></p><p><italic>Specimen examined</italic>: <bold>Kenya</bold>, Coast Province, Malindi District,
Arabuko-Sokoke National Park, 6 Nov. 2006, G.K. Mugambi. Deposited as BPI
879774, <bold>holotype</bold> [formerly, GKM L166A (EA)].</p><p><italic>Notes</italic>: <italic>Gloniopsis arciformis</italic> is represented by a single
specimen (BPI 879774) of only ∼30 fruitbodies in the protected crevice of
a small piece of decorticated hardwood, collected in Arabuko-Sokoke National
Park, Malindi District, Kenya. Although the material is sparse, it does permit
the description of a new species on account of the highly unusual arcuate
dictyospores. <italic>Gloniopsis arciformis</italic> resides in Clade D, and is
phylogenetically closely associated with two other species of
<italic>Gloniopsis</italic> (<italic>Gp. praelonga</italic> and <italic>Gp. subrugosa</italic>), as
well as with an additional new species described below.</p><p><italic><bold>Gloniopsis kenyensis</bold></italic> E.W.A. Boehm, G.K. Mugambi, S.M.
Huhndorf & C.L. Schoch, <bold>sp. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515359&link_type=mb">MB515359</ext-link>.
<xref ref-type="fig" rid="fig7">Fig. 7I–M</xref>.</p><p><italic>Etymology</italic>: From the Latin <italic>-ensis</italic> to denote origin, from
Kenya.</p><p>Hysterothecia navicularia, carbonacea, recta vel flexuosa, utrinque obtusa,
dense aggregata, erumpentia, ad latera inconspicue striata vel levia,
(0.5–)1–3 mm longa, 250–350 μm lata, 250–350 μm
alta. Peridium prope basim ad 100 μm crassum, bivel tristratosum, stratum
internum compressum, hyalinum, strata exteriora densiora et fusca.
Pseudoparaphyses cellulares, septatae, 1–1.5 μm latae, sursum ramosae
et anastomosantes, epithecium pigmentatum ascos obtegens formantes. Asci
cylindrici vel clavati, bitunicati, 60–80 x 12–16 μm,
ascosporas irregulariter biseriatas continentes. Ascosporae dictyoseptatae,
pigmentatae, obovoideae, tenuitunicatae, fragiles, polis asymmetricis: apice
obtuso, ad basim acuminatae vel nonnumquam protrudentes, 3(–4)-septatae,
1–2 septis verticalibus, utrinque saepe septis obliquis divisae, ad
septa vix constrictae, iuvenes guttulis repletae,
(12–)15–18(–19) x 5–7(–8) μm.</p><p><italic>Hysterothecia</italic> navicular, carbonaceous, straight to flexuous, with
obtuse ends, densely aggregated, erumpent, slightly striated laterally to
smooth, (0.5–)1–3 mm long, 250–350 μm wide, by
250–350 μm high. <italic>Peridium</italic> up to 100 μm thick at base,
composed of two to three layers, the inner thin, compressed and hyaline, the
outer two progressively denser, and darkly pigmented.
<italic>Pseudoparaphyses</italic> cellular, septate, 1–1.5 μm wide, branched,
anastomosed distally, forming a pigmented epithecium above the asci.
<italic>Asci</italic> cylindrical to clavate, bitunicate, 60–80 x 12–16
μm (n = 5), ascospores irregularly biseriate. <italic>Ascospores</italic> pigmented
dictyospores, in outline obovoid, thin-walled, very fragile, spore apices
asymmetric, the upper obtuse, the lower acuminate and sometimes drawn out,
3(–4[rarely])-septate, with 1–2 vertical septa, often with oblique
septa in end cell, hardly constricted at the septa, highly gutulate when
young, (12–)15–18(–19) x 5–7(–8) μm (n = 14).
Known from only one collection, from Kenya, East Africa.</p><p><italic>Specimen examined</italic>: <bold>Kenya</bold>, Coast Province, Malindi District,
Arabuko-Sokoke National Park, 6 Apr. 2005, G.K. Mugambi. Deposited as BPI
879775, <bold>holotype</bold>; GKM 1010 (EA), <bold>paratype</bold>.</p><p><italic>Notes</italic>: Molecular data indicate that both <italic>Gp. kenyensis</italic>and <italic>Gp. arciformis</italic> are closely associated, adjacent to <italic>Gp.
praelonga</italic> and <italic>Gp. subrugosa</italic> in Clade D. The spores of all four
taxa, however, are different, and thus their association would not have been
predicted based on traditional morphology. The spores of <italic>Gp.
kenyensis</italic> do bear a close resemblance, however, to those of <italic>Hb.
mori</italic>. Both have predominantly 3-septate, thin-walled, pigmented
dictyospores, with 1–2 vertical septa, often with oblique septa in the
end cell. They can be differentiated on spore size:
(12–)14–22(–26) x (5–)7–10(–11) μm for
<italic>Hb. mori, versus</italic> (12–)15–18(–19) x
5–7(–8) μm for <italic>Gp. kenyensis</italic>. The spores of <italic>Hb.
mori</italic> are usually longer and wider, and also show prominent septal
constrictions, especially when fresh and hydrated. Additionally, <italic>Gp.
kenyensis</italic> is highly guttulate when young, where this is rarely observed
in <italic>Hb. mori</italic>. Molecular data indicate that they are not related.</p><p>To summarise, molecular data have necessitated the break up of the genus
<italic>Hysterographium</italic>, because the type, <italic>Hg. fraxini</italic>, no longer
resides within the <italic>Hysteriaceae</italic>(<xref ref-type="bibr" rid="ref18">Boehm <italic>et al.</italic> 2009</xref>).
This break up has resulted in: (1) the new genus <italic>Hysterobrevium</italic>,
which includes both species with hyaline dictyospores, previously classified
as <italic>Gloniopsis</italic> (<italic>Hb. constrictum</italic> and <italic>Hb. smilacis</italic>),
and species with pigmented dictyospores, previously classified as
<italic>Hysterographium</italic> (<italic>Hb. mori</italic>) in Clade A; (2) the inclusion in
<italic>Gloniopsis</italic> of both hyaline (<italic>Gp. praelonga</italic>) and pigmented
(<italic>Gp. subrugosa, Gp. arciformis, Gp. kenyensis</italic>) dictyospores in Clade
D; (3) the inclusion in <italic>Oedohysterium</italic> of pigmented dictyospored
species previously classified in <italic>Hysterographium</italic> (<italic>Od.
pulchrum</italic>), also in Clade D; and, lastly, (4) the removal of
<italic>Hysterographium</italic>, with the type <italic>Hg. fraxini</italic>, from the
<italic>Hysteriaceae</italic>, currently placed as <italic>Pleosporomycetidae gen.
incertae sedis</italic>. As the taxonomy of <italic>Hysterographium,
Hysterobrevium</italic> and <italic>Gloniopsis</italic> is currently in flux, we chose to
provide the following dichotomous key, whereby all hysteriaceous fungi,
bearing transversely and longitudinally septate dictyospores, whether
pigmented or hyaline, are identified together, with the caveat that unrelated
taxa share the same key.</p></sec><sec><title>Key to the species of <italic>Hysterographium, Hysterobrevium</italic> and
<italic>Gloniopsis</italic></title><p><list list-type="simple"><list-item><p>1. Dictyospores, usually shorter than 25
μm.................................................................................................................................................
2
1. Dictyospores mostly longer than 25
μm....................................................................................................................................................
6</p></list-item><list-item><p>2. Dictyospores pigmented, thin-walled, fragile, pronouncedly arcuate or
bent, 3–5(–7)-septate, with 1–2(–3) vertical septa,
which are mostly associated with the mid-cells, these much larger and swollen
than the end-cells, no septal constrictions, (10–)12–18(–22)
x 6–10 μm;
Kenya............................................................................................
<italic><bold>Gp. arciformis</bold></italic>2. Not with the above combination of
characters..........................................................................................................................................
3</p></list-item><list-item><p>3. Dictyospores hyaline at
maturity...............................................................................................................................................................
4
3. Dictyospores pigmented at
maturity..........................................................................................................................................................
5</p></list-item><list-item><p>4. Dictyospores highly symmetric in outline and septation, with thickened
walls, gelatinous sheath present when young, absent at maturity,
(1–)3(–4)-septate, with 1(–2) vertical septa, that may pass
through one to two cells; (11–)13–20(–23) x 5–12
μm; Japan, New
Zealand.........................................................................................................
<italic><bold>Hb. constrictum</bold></italic>4. Dictyospores asymmetric, with acuminate
ends, with a gelatinous sheath when young, mostly 3–5(–9)-septate
and with 1(–3) vertical septa, passing through multiple mid-cells,
prominently constricted at the median septum, sometimes constriced at multiple
septa, (13–)15–26(–31) x (4–)5–9(–10)
μm; cosmopolitan.............................................. <italic><bold>Hb.
smilacis</bold></italic></p></list-item><list-item><p>5. Dictyospores thin-walled, obovoid, with obtuse ends,
3–(5–7)-septate, with 1–2(–3) vertical septa, usually
associated with mid-cells, but occasionally present obliquely in end-cells,
constricted at the median septum, sometimes at additional septa,
(12–)14–22(–26) x (5–)7–10(–11) μm;
cosmopolitan.................................................................
<italic><bold>Hb. mori</bold></italic>5. Dictyospores thin-walled, very fragile,
obovoid, 3[–4(rarely)]-septate, with 1–2 vertical septa, highly
gutulate when young, spore apices asymmetric, the upper obtuse, the lower
acuminate and sometimes drawn out, often with oblique septa in end cell(s),
hardly constricted at the septa, measuring (12–)15–18(–19) x
5–7(–8) μm; Kenya.......................................
<italic><bold>Gp. kenyensis</bold></italic></p></list-item><list-item><p>6. Red pigment present in hamathecium and/or centrum; dictyospores
pigmented.....................................................................................
7
6. No red pigment present, spores pigmented or
hyaline.............................................................................................................................
8</p></list-item><list-item><p>7. Dictyospores, 22–25(–27) x 5–6 μm, with (5–)6
transverse and 1 vertical septum in either cell or both cells adjacent to the
primary septum; typically with red pigment in the hamathecium; neotropical
(Costa Rica)....................... <italic><bold>Od. pulchrum</bold>Note</italic>:
<italic>Od. pulchrum</italic> is accommodated in the genus <italic>Oedohysterium</italic> and
is present in both keys.
7. Dictyospores 25–28 x 11–13 μm,
with 5–6 transverse and mostly one longitudinal septum; hamathecium
brick-red; on <italic>Acacia</italic> thorns, South
Africa......................................................................................................
<italic><bold>Hg. spinicola</bold></italic></p></list-item><list-item><p>8. Dictyospores hyaline or turning brown
tardily...........................................................................................................................................
9
8. Dictyospores pigmented in the
ascus.....................................................................................................................................................
10</p></list-item><list-item><p>9. Dictyospores hyaline turning yellow in age, obovoid, ends usually
obtuse, 5–7(–10)-septate, with 2–3 longitudinal septa,
constricted at the median and often other septa, gelatinous sheath when young,
(16–)20–32(–34) x (6–)9–12(–15) μm;
cosmopolitan.........................................................................................................
<italic><bold>Gp. praelonga</bold></italic>9. Ascospores irregularly biseriate,
ellipsoid, hyaline but becoming brown tardily, with the upper half generally
wider than the lower half, sometimes surrounded by a gelatinous sheath, with
7–13 transverse and 1–3 longitudinal septa, constricted at the
median transverse septum; 25–49 x 8–17 μm;
Japan................................................................................................................................................
<italic><bold>Gp. macrospora</bold></italic></p></list-item><list-item><p>10. Dictyospores usually less than 38 μm
long............................................................................................................................................
11
10. Dictyospores 30–80 μm
long..................................................................................................................................................................
12</p></list-item><list-item><p>11. Dictyospores (22–)25–34(–45) x
(6–)8–12(–17) μm, mostly with 7–11 transverse and
1–2 vertical septa;
cosmopolitan.....................................................................................................................................................................
<italic><bold>Gp. subrugosa</bold></italic>11. Dictyospores 26–38 x 10–15
μm, with 6–13 transverse and 1–3 vertical septa, obovoid, ends
obtuse; Japan....................... <italic><bold>Hg. minus</bold></italic></p></list-item><list-item><p>12. Dictyospores (25–)30–45(–51) x
(10–)12–15(–22) μm, with 7–9 transverse and
2–3 vertical septa, obovoid, ends obtuse;
cosmopolitan.........................................................................................................................................
<italic><bold>Hg. fraxini</bold>Note</italic>: <italic>Hysterographium fraxini</italic>, the
type species for the genus <italic>Hysterographium</italic>, lies outside of the
<italic>Hysteriaceae</italic>, as <italic>Pleosporomycetidae incertae sedis</italic>(<xref ref-type="bibr" rid="ref18">Boehm <italic>et al.</italic>2009</xref>).
12. Ascospore outline ellipsoid, fusoid, ends slightly
acuminate, (30–)40–65(–80) x
(8–)10–18(–19) μm, with 7–15 transverse and
1–3 vertical septa;
cosmopolitan..............................................................................................
<italic><bold>Hg. flexuosum</bold></italic></p></list-item></list></p><p><fig position="float" id="fig8"><label>Fig. 8.</label><caption><p>The genus <italic>Psiloglonium</italic> (Clade B). A–D. <italic>Psiloglonium
simulans</italic> [ANM 1557 (BPI 879803), U.S.A.]; E–H. <italic>Psiloglonium
clavisporum</italic> [GKM 344A (BPI 879801), Kenya]; I–M. <italic>Psiloglonium
lineare</italic> [ANM 117 (ILLS), U.S.A.; not incl.]; N–Q. <italic>Psiloglonium
araucanum</italic> [ANM 42 (ILLS), U.S.A.; not incl.]. Scale bar (habitat) = 500
μm; Scale bar (spores and asci) = 10 μm.</p></caption><graphic xlink:href="49fig8"></graphic></fig></p><p><italic><bold>Psiloglonium</bold></italic> Höhn.<bold>,</bold> Ann. Mycol. 16: 145.
1918.</p><p>A discussion of the genus <italic>Psiloglonium</italic>(<xref ref-type="bibr" rid="ref41">von Höhnel 1918</xref>; Petrak
<xref ref-type="bibr" rid="ref86">1923a</xref>,
<xref ref-type="bibr" rid="ref87">b</xref>) by necessity must begin
with the genus <italic>Glonium</italic>. This is because Zogg
(<xref ref-type="bibr" rid="ref122">1962</xref>) synonymised a number
of species under the genus <italic>Glonium</italic> that were originally classified in
<italic>Psiloglonium</italic> by von Höhnel
(<xref ref-type="bibr" rid="ref41">1918</xref>) and Petrak
(<xref ref-type="bibr" rid="ref86">1923a</xref>,
<xref ref-type="bibr" rid="ref87">b</xref>). Both <italic>Psiloglonium</italic>and <italic>Glonium</italic> possess hyaline to yellow didymospores, somewhat
constricted at the septum, with obtuse or acuminate ends, typically with cells
unequal in size, borne in hysterothecia.</p><p>Von Höhnel (<xref ref-type="bibr" rid="ref41">1918</xref>) was
the first to view the genus <italic>Glonium</italic> as comprised of two distinct
morphological types, and stressed the importance of subicula, using it to
divide the genus, at first, into two subgenera, <italic>Glonium</italic> and
<italic>Psiloglonium</italic>, and, further in the same article, into two separate
genera, with or without subicula, respectively. Petrak
(<xref ref-type="bibr" rid="ref86">1923a</xref>) recognised that von
Höhnel (<xref ref-type="bibr" rid="ref41">1918</xref>) had
established the genus <italic>Psiloglonium</italic>, both at sub-generic and generic
rank, but it was Petrak
(<xref ref-type="bibr" rid="ref86">1923a</xref>) who explicitly
designated the type species for <italic>Psiloglonium</italic> as <italic>P. lineare</italic>(<xref ref-type="fig" rid="fig8">Fig. 8I–M</xref>), retaining
<italic>G. stellatum</italic> as the type species for the genus <italic>Glonium sensu</italic>von Höhnel (<xref ref-type="bibr" rid="ref41">1918</xref>). Petrak
(<xref ref-type="bibr" rid="ref86">1923a</xref>,
<xref ref-type="bibr" rid="ref87">b</xref>) eventually placed a number
of species in <italic>Psiloglonium</italic>, all subsequently transferred to
<italic>Glonium</italic> by Zogg
(<xref ref-type="bibr" rid="ref122">1962</xref>). Müller & von
Arx (<xref ref-type="bibr" rid="ref82">1950</xref>) originally accepted
the genus <italic>Psiloglonium</italic>, but later reduced it to a synonym of
<italic>Glonium</italic> (<xref ref-type="bibr" rid="ref3">von Arx & Müller
1975</xref>). Lohman
(<xref ref-type="bibr" rid="ref58">1933a</xref>,
<xref ref-type="bibr" rid="ref61">1937</xref>) also did not support
<italic>Psiloglonium</italic>, based on the observation that similar anamorphs were
shared between species of the two subgenera. Barr
(<xref ref-type="bibr" rid="ref7">1987</xref>), was the only modern
author to retain the genus <italic>Psiloglonium</italic>, as distinct from the
subiculate <italic>Glonium</italic>.</p><p>Although von Höhnel
(<xref ref-type="bibr" rid="ref41">1918</xref>) and Petrak
(<xref ref-type="bibr" rid="ref86">1923a</xref>,
<xref ref-type="bibr" rid="ref87">b</xref>) both stressed the
importance of subicula as a major morphological distinction between
<italic>Psiloglonium</italic> and <italic>Glonium</italic>, Zogg
(<xref ref-type="bibr" rid="ref122">1962</xref>) noted that some
species previously classified as <italic>Psiloglonium</italic> by Petrak
(<xref ref-type="bibr" rid="ref86">1923a</xref>) do in fact possess
subicula on occasion (<italic>e.g., P. lineare</italic>). Zogg
(<xref ref-type="bibr" rid="ref122">1962</xref>) further noted an
additional two species that were occasionally associated with subicula, namely
<italic>G. pusillum</italic> and <italic>G. graphicum</italic>, stating: “<italic>...ohne
Subiculum oder auf ziemlich deutlichem Subiculum sitzend...</italic>” Hence,
Zogg (<xref ref-type="bibr" rid="ref122">1962</xref>) considered
subicula not to be a synapomorphic character state, and transferred those
species previously classified by Petrak
(<xref ref-type="bibr" rid="ref86">1923a</xref>,
<xref ref-type="bibr" rid="ref87">b</xref>) in <italic>Psiloglonium</italic>(<italic>e.g., P. lineare, P. microspermum, P. ruthenicum</italic>, and <italic>P.
finkii</italic>) to the genus <italic>Glonium</italic>.</p><p>Although Zogg (<xref ref-type="bibr" rid="ref122">1962</xref>) did
not support <italic>Psiloglonium</italic>, he did in fact recognise three distinct
morphological forms within his concept of <italic>Glonium</italic>, two of which
(Types I and II) we incorporate in <italic>Psiloglonium</italic>, the third (Type III)
forming the basis for the <italic>Gloniaceae</italic>(<xref ref-type="bibr" rid="ref18">Boehm <italic>et al.</italic> 2009</xref>).
Zogg (<xref ref-type="bibr" rid="ref122">1962</xref>) arranged the
species of <italic>Glonium</italic> based on (1) didymospore shape: spore apices
obovoid to rounded (Type I) <italic>versus</italic> spores fusiform with acuminate
apices (Type II and III); and (2) the degree of complexity surrounding the
architecture of the hysterothecia, simple, linear, solitary to gregarious
(Types I, II) <italic>versus</italic> complex bifurcating, laterally anastomosing to
form flabelliform pseudostellate composites, sometimes associated with a thin
stromal crust (Type III). Thus, the genus <italic>Glonium sensu</italic> Zogg
(<xref ref-type="bibr" rid="ref122">1962</xref>) was comprised of two
groups of species, one with obovoid to rounded spores apices borne in regular
hysterothecia (Type I) <italic>versus</italic> those with acuminate spore apices borne
in complex bifurcating or modified hysterothecia (Type III). Species belonging
to Type II possess fruitbodies of Type I, but spores of Type III; the
assumption was that they constituted an intermediate, perhaps transitional,
morphological group. This, then, de-emphasised the presence or absence of
subicula <italic>per se</italic>, as stressed by von Höhnel
(<xref ref-type="bibr" rid="ref41">1918</xref>) and Petrak
(<xref ref-type="bibr" rid="ref86">1923a</xref>,
<xref ref-type="bibr" rid="ref87">b</xref>). Nevertheless, Zogg
(<xref ref-type="bibr" rid="ref122">1962</xref>) maintained all three
types within the genus <italic>Glonium</italic>. Molecular data presented here (see
below), indicate that Types I & II are closely related, with Type III
forming a distant clade in the <italic>Gloniaceae</italic>(<xref ref-type="bibr" rid="ref18">Boehm <italic>et al.</italic>2009</xref>).</p><p><bold>Type I:</bold> This type is characterised by hysterothecia that may be
solitary to gregarious, erumpent to entirely superficial, navicular to linear
to highly flexuous, even triradiate, sometimes arranged in parallel
orientation and confluent linearly to some degree, but never dichotomously
branched, or associated with a stromal crust, as found in the
<italic>Gloniaceae</italic> (Type III). These species correspond to <italic>Psiloglonium
sensu</italic> von Höhnel
(<xref ref-type="bibr" rid="ref41">1918</xref>). Here, the didymospores
are relatively small, hyaline, and have at least one, if not both ends,
obovoid to obtuse (Type I), rather than acuminate (Types II and III). Zogg
(<xref ref-type="bibr" rid="ref122">1962</xref>) recognised five
species, listed here by increasing ascospore length: <italic>Glonium abbreviatum,
G. pusillum, G. lineare, G. chambianum</italic>, and <italic>G. curtisii.</italic> Barr
(<xref ref-type="bibr" rid="ref4">1975</xref>) transferred the last
species to <italic>Ostreichnion</italic>, as <italic>O. curtisii</italic> in the
<italic>Mytilinidiaceae</italic>, since transferred to the <italic>Hysteriaceae</italic>(<xref ref-type="bibr" rid="ref18">Boehm <italic>et al.</italic> 2009</xref>).
A sixth species, <italic>G. finkii</italic>, was included by Zogg
(<xref ref-type="bibr" rid="ref122">1962</xref>), based on ascospore
shape, but placed apart in the key due to the unusual arrangement of the
ascospores within the upper part of the ascus
(<xref ref-type="bibr" rid="ref61">Lohman 1937</xref>).</p><p><italic>Psiloglonium lineare</italic> was previously reinstated within the
<italic>Hysteriaceae</italic>, listing <italic>G. lineare</italic> as a synonym
(<xref ref-type="bibr" rid="ref18">Boehm <italic>et al.</italic> 2009</xref>).
Here we also reinstate <italic>Psiloglonium finkii</italic>. An additional two species
are included in Type I, namely <italic>G. simulans</italic> and <italic>G.
clavisporum</italic>, synonymised by Zogg
(<xref ref-type="bibr" rid="ref122">1962</xref>) under <italic>G.
lineare</italic>, but earlier recognised by Lohman
(<xref ref-type="bibr" rid="ref56">1932a</xref>,
<xref ref-type="bibr" rid="ref61">1937</xref>) to be distinct from
<italic>G. lineare</italic>. Boehm <italic>et al.</italic>(<xref ref-type="bibr" rid="ref18">2009</xref>) proposed new
combinations for these taxa, based on morphological as well as molecular data,
as <italic>P. simulans</italic> (<xref ref-type="fig" rid="fig8">Fig.
8A–D</xref>) and <italic>P. clavisporum</italic>(<xref ref-type="fig" rid="fig8">Fig. 8E–H</xref>). To these
species can also be added <italic>G. sasicola</italic> from Japan, the first report of
a gelatinous sheath in the genus (<xref ref-type="bibr" rid="ref1">Amano
1983</xref>). In this same publication Amano
(<xref ref-type="bibr" rid="ref1">1983</xref>) proposed an additional
new species, <italic>G. macrosporum</italic>, also from Japan. The spore measurements
were given as 13.1–16.8 x 4–5.6 μm, nearly identical to those
of <italic>P. simulans</italic> at (10–)14–16(–18) x
(4.5–)5–6 μm (<xref ref-type="bibr" rid="ref61">Lohman
1937</xref>). Moreover, the illustrations given by Amano
(<xref ref-type="bibr" rid="ref1">1983</xref>) match closely those
given by Lohman (<xref ref-type="bibr" rid="ref56">1932a</xref>) for
<italic>P. simulans</italic>. We therefore synonymise <italic>G. macrosporum</italic> under
<italic>P. simulans</italic>.</p><p>More recently, Lorenzo & Messuti
(<xref ref-type="bibr" rid="ref62">1998</xref>), in a reappraisal of
the type specimens collected by Spegazzini and Hennings from Argentina and
Chile, have reinstated <italic>Glonium costesii</italic>. In a later publication,
Messuti & Lorenzo (<xref ref-type="bibr" rid="ref77">2007</xref>)
synonymised <italic>G. costesii</italic> under the earlier epithet <italic>G.
ephedrae</italic>. With spore measurements of 26–35 x 8–15 μm,
<italic>G. ephedrae</italic> possesses the largest spores in Type I. In the same
publication, Messuti & Lorenzo
(<xref ref-type="bibr" rid="ref77">2007</xref>) also accepted two
additional species, <italic>G. chilense</italic> and <italic>G. uspallatense</italic>,
previously considered by Zogg
(<xref ref-type="bibr" rid="ref122">1962</xref>) to be doubtful
species. The spores of <italic>G. chilense</italic> measure 15–16 x
(5–)7–8 μm, which places it very close to <italic>P. lineare</italic>,
the latter with slightly smaller spores, (10–) 12–14(–18) x
(4–)5–7(–8) μm (<xref ref-type="bibr" rid="ref122">Zogg
1962</xref>). However, <italic>G. chilense</italic> has almost identical
ascomatal and spore measurements as <italic>P. simulans</italic>, given above. We
therefore synonymise <italic>G. chilense</italic> with the earlier name <italic>G.
simulans</italic>, as <italic>P. simulans</italic>. For <italic>G. uspallatense</italic>, Messuti
& Lorenzo (<xref ref-type="bibr" rid="ref77">2007</xref>) gave
spore measurements of 18–24 x 10–12 μm, intermediate between
<italic>G. chambianum</italic>, (14–)16–18(–21) x
(6–)8–9(–10) μm
(<xref ref-type="bibr" rid="ref122">Zogg 1962</xref>), and <italic>G.
sasicola</italic>, 25–32 x 5–8 μm
(<xref ref-type="bibr" rid="ref1">Amano 1983</xref>).</p><p>Recently, Mugambi & Huhndorf
(<xref ref-type="bibr" rid="ref81">2009</xref>) proposed a new genus,
<italic>Anteaglonium</italic>, outside of the <italic>Hysteriales</italic> but within the
<italic>Pleosporales</italic>, to accommodate <italic>A. abbreviatum</italic>(<xref ref-type="fig" rid="fig9">Fig. 9A–E</xref>), <italic>A.
globosum</italic> (<xref ref-type="fig" rid="fig9">Fig.
9F–I</xref>), <italic>A. parvulum</italic>(<xref ref-type="fig" rid="fig9">Fig. 9J–M</xref>), and <italic>A.
latirostrum</italic> (<xref ref-type="fig" rid="fig9">Fig.
9N–R</xref>). The first three species are characterised by hyaline
didymospores that belong to Type I, as defined by Zogg
(<xref ref-type="bibr" rid="ref122">1962</xref>), and are less than 8
μm in length. The fourth species, <italic>A. latirostrum</italic>, belongs to Type
II (see below), with longer spores. Although phylogenetically unrelated to
<italic>Psiloglonium</italic>, these species share a similar morphology and thus are
included in the key below.</p><p><fig position="float" id="fig9"><label>Fig. 9.</label><caption><p>The genus <italic>Anteaglonium</italic> (<italic>Pleosporales</italic>). A–E.
<italic>Anteaglonium abbreviatum</italic> [ANM 37 (ILLS), U.S.A.; not incl.];
F–I. <italic>Anteaglonium globosum</italic> [ANM 925.2 (ILLS), U.S.A.];
J–M. <italic>Anteaglonium parvulum</italic> [GKM 219N (EA), Kenya; not incl.];
N–R. <italic>Anteaglonium latirostrum</italic> [GKM L100N.2 (EA), Kenya]. Scale
bar (habitat) = 500 μm; Scale bar (spores and asci) = 5 μm.</p></caption><graphic xlink:href="49fig9"></graphic></fig></p><p><bold>Type II:</bold> This type is characterised by relatively large
didymospores, distinctly fusoid in outline, prominently constricted at the
septum, and with acuminate apices. Zogg
(<xref ref-type="bibr" rid="ref122">1962</xref>) recognised two
species, namely <italic>Glonium caucasicum</italic> and the much larger-spored,
neotropical <italic>G. hysterinum</italic>, to which can be added the newly described
<italic>G. colihuae</italic>, on <italic>Chusquea culeou</italic> from Argentina
(<xref ref-type="bibr" rid="ref62">Lorenzo & Messuti 1998</xref>).
<italic>Glonium caucasicum</italic> has recently been synonymised under the earlier
name <italic>G. araucanum</italic> by Messuti & Lorenzo
(<xref ref-type="bibr" rid="ref77">2007</xref>), based on a comparison
of the type specimen of <italic>G. caucasicum</italic> to Spegazini's earlier type of
<italic>G. araucanum</italic> from Chile.</p><p><bold>Type III:</bold> This type corresponds to von Höhnel's
(<xref ref-type="bibr" rid="ref41">1918</xref>) and Petrak's
(<xref ref-type="bibr" rid="ref86">1923a</xref>,
<xref ref-type="bibr" rid="ref87">b</xref>) circumscription of the
genus <italic>Glonium</italic>, and includes species with fusiform spores, with
acuminate apices, typically producing complex laterally anastomosing
hysterothecia, forming stellate composites, usually with prominent subicula,
with or without stroma. Zogg
(<xref ref-type="bibr" rid="ref122">1962</xref>) included the type,
<italic>G. stellatum</italic> (<xref ref-type="fig" rid="fig12">Fig.
12A–E</xref>), <italic>G. compactum</italic>, and <italic>G. graphicum</italic>,
the later sometimes variably associated with subicula. Zogg
(<xref ref-type="bibr" rid="ref122">1962</xref>) also stated that
<italic>G. compactum</italic> possesses a subiculum, much like <italic>G. stellatum</italic>,
and with similar spore size, but whereas hysterothecia in <italic>G.
stellatum</italic> are merely seated on the subiculum, in <italic>G. compactum</italic>the hysterothecia are embedded in and arise from a thin stromal crust, which
is itself seated on subicula. Recently, a fourth species was added, based on
molecular evidence (<xref ref-type="bibr" rid="ref18">Boehm <italic>et al.</italic>2009</xref>), namely <italic>G. circumserpens</italic>(<xref ref-type="fig" rid="fig12">Fig. 12F–H</xref>), from
Tasmania (<xref ref-type="bibr" rid="ref43">Kantvilas & Coppins
1997</xref>).</p><p>Sequence data presented here (<xref ref-type="fig" rid="fig1">Fig.
1</xref>) and elsewhere (<xref ref-type="bibr" rid="ref18">Boehm <italic>et
al.</italic> 2009</xref>, <xref ref-type="bibr" rid="ref81">Mugambi &
Huhndorf 2009</xref>), clearly indicate that the genus <italic>Glonium
sensu</italic> Zogg (<xref ref-type="bibr" rid="ref122">1962</xref>)
actually comprises three entirely unrelated lineages within the
<italic>Pleosporomycetidae</italic>, one within the <italic>Hysteriaceae</italic> and two
forming clades outside of the family. The first lineage corresponds to
<italic>Psiloglonium sensu</italic> von Höhnel
(<xref ref-type="bibr" rid="ref41">1918</xref>), and forms a highly
supported monophyletic clade in this study (Clade B in
<xref ref-type="fig" rid="fig1">Fig. 1</xref>). This clade includes:
<italic>Psiloglonium clavisporum</italic>, with four single-ascospore isolates from
New Jersey, the United States
(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123338&link_type=cbs">CBS 123338</ext-link> / BPI
878726, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123339&link_type=cbs">CBS 123339</ext-link>/ BPI 878727, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123340&link_type=cbs">CBS
123340</ext-link> / BPI 878728 and
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123341&link_type=cbs">CBS 123341</ext-link> / BPI
878729), and two from Kenya (GKM 344A / BPI 879801, GKM L172A in EA), <italic>P.
simulans</italic>, with two isolates from the United States, one from Michigan
(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=206.34&link_type=cbs">CBS 206.34</ext-link>),
deposited in 1934 by Lohman, and a more recent collection from Tennessee (ANM
1557 / BPI 879803), and, lastly, <italic>P. araucanum</italic>, with three isolates
from South Africa, two from Kirstenbosch
(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=112412&link_type=cbs">CBS 112412</ext-link> / PREM
57570, CMW 18760/ PREM 57569) and one from Jonkershoek (CMW 17941 / PREM
575566). <italic>Psiloglonium clavisporum</italic> and <italic>P. simulans</italic> belong to
Type I, whereas <italic>P. araucanum</italic> belongs to Type II. Both are
phylogenetically related and reside in Clade B
(<xref ref-type="fig" rid="fig1">Fig. 1</xref>). Recently, a second
lineage has been shown to be associated with the <italic>Pleosporales</italic>, now
accommodated in the new genus <italic>Anteaglonium</italic>(<xref ref-type="bibr" rid="ref81">Mugambi & Huhndorf 2009</xref>),
for which we include six accessions representing four species
(<xref ref-type="table" rid="tbl1">Table 1</xref>). The third lineage
corresponds to <italic>Glonium</italic> (Type III), in the <italic>Gloniaceae</italic>(<xref ref-type="bibr" rid="ref18">Boehm <italic>et al.</italic> 2009</xref>),
for which we have included four isolates, representing two species
(<xref ref-type="table" rid="tbl1">Table 1</xref>). We treat here all
species of <italic>Glonium sensu</italic> Zogg
(<xref ref-type="bibr" rid="ref122">1962</xref>), belonging to Types I
and II, outside of <italic>Anteaglonium</italic>, as belonging to
<italic>Psiloglonium</italic>. Since the generic name <italic>Glonium</italic> is reserved for
species in the <italic>Gloniaceae</italic> (<xref ref-type="bibr" rid="ref18">Boehm
<italic>et al.</italic> 2009</xref>), we propose eight new combinations for the
genus <italic>Psiloglonium</italic>.</p><p><italic><bold>Psiloglonium pusillum</bold></italic> (H. Zogg) E.W.A. Boehm & C.L.
Schoch, <italic><bold>comb. nov.</bold></italic> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515327&link_type=mb">MB515327</ext-link>.</p><p><italic>Basionym</italic>: <italic>Glonium pusillum</italic> H. Zogg, Beitr. Kryptogamenfl.
Schweiz. 11(3): 62. 1962.</p><p><italic>Notes</italic>: Zogg
(<xref ref-type="bibr" rid="ref122">1962</xref>) described this species
as <italic>G. pusillum</italic> from <italic>Juniperus phoenicea</italic> and <italic>Pinus
sylvestris</italic> from Southern France, noting that it was quite rare. Zogg
(<xref ref-type="bibr" rid="ref122">1962</xref>) stated that this
species may or may not be associated with a subiculum, and hence was one of
the factors behind his transfer of Petrak's
(<xref ref-type="bibr" rid="ref86">1923a</xref>,
<xref ref-type="bibr" rid="ref87">b</xref>) <italic>Psiloglonium</italic>species to <italic>Glonium. Psiloglonium pusillum</italic> has ascospores only
slightly larger than those of <italic>P. abbreviatum</italic>, measuring
(9–)10–12(–13) x 4–5(–6) μm. Lee & Crous
(<xref ref-type="bibr" rid="ref49">2003</xref>) also identified this
fungus from <italic>Proteaceae</italic> and <italic>Restionaceae</italic> in South Africa, and
Sivanesan & Hsieh (<xref ref-type="bibr" rid="ref104">1989</xref>)
reported it from Taiwan. It has also been found in North America (Boehm,
unpubl. data).</p><p><italic><bold>Psiloglonium chambianum</bold></italic> (Guyot) E.W.A. Boehm & C.L.
Schoch, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515320&link_type=mb">MB515320</ext-link>.</p><p><italic>Basionym</italic>: <italic>Glonium chambianum</italic> Guyot, Ann. Serv. Bot.
Tunisie 28: 90. 1955.</p><p><italic>Notes</italic>: Originally from North Africa, on <italic>Lonicera implexa</italic>(<italic>Caprifoliaceae</italic>), the fungus has since been reported from the
<italic>Proteaceae</italic> in South Africa (<xref ref-type="bibr" rid="ref49">Lee
& Crous 2003</xref>) and Europe. Zogg
(<xref ref-type="bibr" rid="ref122">1962</xref>) gave the spore
measurements for <italic>G. chambianum</italic> as (14–)16–18(–21) x
(6–)8–9(–10) μm, whereas Lee & Crous
(<xref ref-type="bibr" rid="ref49">2003</xref>) gave slightly larger
measurements, (18–)20–21(–23) x
(4–)5–6(–7) μm. Spores ellipsoid to oblong, with upper
cell broader than the lower, and with an obovoid, obtuse apex.
<italic>Psiloglonium chambianum</italic> possesses larger spores than <italic>P. lineare,
P. simulans</italic>, and <italic>P. clavisporum</italic>, but smaller than <italic>P.
uspallatense</italic>.</p><p><italic><bold>Psiloglonium uspallatense</bold></italic> (Speg.) E.W.A. Boehm & C.L.
Schoch, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515321&link_type=mb">MB515321</ext-link>.</p><p><italic>Basionym</italic>: <italic>Glonium uspallatense</italic> Speg., Anales Mus. Nac.
Hist. Nat. Buenos Aires. 19: 436. 1909.</p><p><italic>Notes</italic>: Zogg
(<xref ref-type="bibr" rid="ref122">1962</xref>) listed the species a
“doubtful”, but Messuti & Lorenzo
(<xref ref-type="bibr" rid="ref77">2007</xref>) reinstated <italic>G.
uspallatense</italic> after locating the original holotype material. They gave the
spore measurements as 18–24 x 10–12 μm, placing it intermediate
between <italic>P. chambianum</italic> and <italic>P. sasicola</italic>.</p><p><italic><bold>Psiloglonium sasicola</bold></italic> (N. Amano) E.W.A. Boehm & C.L.
Schoch, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515322&link_type=mb">MB515322</ext-link>.</p><p><italic>Basionym</italic>: <italic>Glonium sasicola</italic> N. Amano, Trans. Mycol. Soc.
Japan 24: 287. 1983.</p><p><italic>Notes</italic>: Amano (<xref ref-type="bibr" rid="ref1">1983</xref>)
described this species from dead culms of <italic>Sasa</italic> sp.
(<italic>Bambusaceae</italic>) in Japan. The ascospore measurements were given as
25–32 x 5–8 μm, with a rounded apical cell, placing it between
<italic>P. uspallatense</italic> and <italic>P. ephedrae</italic>. Amano
(<xref ref-type="bibr" rid="ref1">1983</xref>) further reported that
ascospores of this species are associated with a gelatinous sheath, previously
not known among these didymospored fungi.</p><p><italic><bold>Psiloglonium ephedrae</bold></italic> (Henn.) E.W.A. Boehm & C.L.
Schoch, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515323&link_type=mb">MB515323</ext-link>.
<italic>Basionym</italic>:
<italic>Glonium ephedrae</italic> Henn., Öfvers. K. Vet. Akad. Förhandl. 2:
328. 1900.</p><p><list list-type="simple"><list-item><p>= <italic>Glonium costesi</italic> Speg., Bol., Acad. Ci., Córdoba 25: 78.
1921.</p></list-item></list></p><p><italic>Notes</italic>: Messuti & Lorenzo
(<xref ref-type="bibr" rid="ref77">2007</xref>) reinstated <italic>G.
ephedrae</italic> with the synonym <italic>G. costesi</italic>, after locating and
comparing original type materials. <italic>Psiloglonium ephedrae</italic> possesses
very large didymospores, measuring 26–35 x 8–15 μm, the upper
cells broadly ovate. It has been collected from <italic>Ephedra andicola</italic>,
and, as <italic>G. costesi</italic>, from <italic>Proustia pyrifolia</italic> in Chile.</p><p><italic><bold>Psiloglonium hysterinum</bold></italic> (Rehm) E.W.A. Boehm & C.L.
Schoch, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515324&link_type=mb">MB515324</ext-link>.</p><p><italic>Basionym</italic>: <italic>Glonium hysterinum</italic> Rehm, Hedwigia 37: 298.
1898.</p><p><italic>Notes</italic>: Rehm (<xref ref-type="bibr" rid="ref89">1898</xref>)
originally described a species of <italic>Glonium</italic> from Southern Brazil with
large fusiform didymospores, prominently constricted at the septum, and with
acuminate spore apices (“<italic>Enden zugespitzt</italic>”). The spore
measurements were given as 45 x 9 μm.</p><p><italic><bold>Psiloglonium colihuae</bold></italic> (Lorenzo & Messuti) E.W.A. Boehm
& C.L. Schoch, <bold>comb. nov.</bold> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515325&link_type=mb">MB515325</ext-link>.</p><p><italic>Basionym</italic>: <italic>Glonium colihuae</italic> Lorenzo & Messuti, Mycol.
Res. 102: 1104. 1998.</p><p><italic>Notes</italic>: Lorenzo & Messuti
(<xref ref-type="bibr" rid="ref62">1998</xref>) described a new species
on culms of <italic>Chusquea culeou</italic> from the Argentine <italic>Nothofagus</italic>rainforests. The spore measurements were given as 30–43 x 4–9.8
μm, and, although the spores are fusiform in outline, they possess
moderately acuminate apices. In comparing this species to other
acuminate-spored species of <italic>Glonium</italic>, the authors noted that the
greatest degree of similarity was with the slightly smaller-spored <italic>G.
caucasicum</italic>.</p><p><italic><bold>Psiloglonium araucanum</bold></italic> (Speg.) E.W.A. Boehm, S. Marincowitz
& C.L. Schoch, <bold>comb. nov</bold><italic>.</italic> MycoBank
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB515326&link_type=mb">MB515326</ext-link>.
<xref ref-type="fig" rid="fig8">Fig.
8N–Q</xref>.
<italic>Basionym</italic>: <italic>Glonium araucanum</italic>Speg., Revista Fac. Agron. Univ. Nac. La Plata 6: 110. 1910.</p><p><list list-type="simple"><list-item><p>= <italic>Gloniella caucasica</italic> Rehm, Vestn. Tiflissk. Bot. Sada 25:12.
1912.</p></list-item><list-item><p>≡ <italic>Glonium caucasicum</italic> (Rehm) H. Zogg, Beitr. Kryptogamenfl.
Schweiz. 11(3): 67. 1962.</p></list-item></list></p><p><italic>Notes</italic>: Messuti & Lorenzo
(<xref ref-type="bibr" rid="ref77">2007</xref>) transferred <italic>Glonium
caucasicum</italic> to <italic>G. araucanum</italic>, after examining the types for both
species. Previously, Zogg
(<xref ref-type="bibr" rid="ref122">1962</xref>) had transferred
<italic>Gloniella caucasica</italic> to <italic>Glonium</italic>. Here we transfer <italic>G.
araucanum</italic> to <italic>Psiloglonium</italic>. This taxon possesses fusiform spores
with highly acuminate apices. Messuti & Lorenzo gave the spore
measurements as 22–28 x 8–10 μm, whereas Zogg
(<xref ref-type="bibr" rid="ref122">1962</xref>) gives them as
(19–)22–25(–27) x (6–) 7–9(–10) μm.
Although originally European in distribution
(<xref ref-type="bibr" rid="ref122">Zogg 1962</xref>), the taxon has
subsequently been collected from South
(<xref ref-type="bibr" rid="ref77">Messuti & Lorenzo 2007</xref>)
and North America (Boehm unpubl. data), and from South Africa
(<xref ref-type="bibr" rid="ref49">Lee & Crous 2003</xref>).</p></sec><sec><title>Key to the species of <italic>Psiloglonium</italic> and
<italic>Anteaglonium</italic></title><p><list list-type="simple"><list-item><p>1. Asci ovoid, +/- cylindrical; ascospores borne in the upper portion of
the ascus, not evenly distributed; ascospores (12–)13–15 x
6–7 μm; Puerto
Rico..............................................................................................................................
<italic><bold>P. finkii</bold></italic>1. Asci typically cylindrical to club-shaped;
ascospores in one row or distichous in the asci, but always regularly arranged
for its full
length.........................................................................................................................................
2</p></list-item><list-item><p>2. Ascospores obovoid, with at least one, often both, ends obtuse,
typically with upper cell larger, +/- constricted at the septum (Type
I).......................................................................................................................................................
3
2. Ascospores fusiform (<italic>i.e.</italic>, spindle-shaped), with both ends
acuminate, usually constricted at the septum (Types II and
III)................. 14</p></list-item><list-item><p>3. Ascospores small, 8 μm or less in length (<italic>Anteaglonium</italic>, in
part)............................................................................................................
4
3. Ascospores longer than 8 μm (<italic>Psiloglonium</italic> Type
I).................................................................................................................................
6</p></list-item><list-item><p>4. Ascospores 6–8 x 2.5–3 μm; hysterothecia with apices
acuminate, but not associated with a darkened crust; no KOH-soluble pigments;
New Zealand, East Africa, North
America...................................................................................
<italic><bold>A. parvulum</bold>Note</italic>: <italic>A. parvulum</italic> lies within the
<italic>Pleosporales</italic> (<xref ref-type="bibr" rid="ref81">Mugambi &
Huhndorf 2009</xref>).
4. Not with the above combination of
characters..........................................................................................................................................
5</p></list-item><list-item><p>5. Ascospores (5–)6–7(–8) x 2–3(–3.5) μm
(as in <italic>A. parvulum</italic>); but hysterothecia with apices truncated, and
associated with a darkened crust (tending to darken the substratum); minute
amounts of soluble pigment in KOH (easily missed); Europe, East Africa, North
America..............................................................................
<italic><bold>A. abbreviatum</bold>Note</italic>: <italic>A. abbreviatum</italic> lies within
the <italic>Pleosporales</italic> (<xref ref-type="bibr" rid="ref81">Mugambi &
Huhndorf 2009</xref>)
5. Ascospores 6–7 x 2–3 μm (as in
<italic>A. parvulum</italic> and <italic>A. abbreviatum</italic>); but hysterothecia globose
with roughened walls, an indistinct slit, and associated with sparse, short
subicula, and also with short tomentum on the walls of the ascomata; like
<italic>A. abbreviatum</italic> also associated with a darkened crust on substrate;
producing a strong green soluble pigment in KOH; eastern and mid-western North
America...............................................................................................................................
<italic><bold>A. globosum</bold>Note</italic>: <italic>A. globosum</italic> lies within the
<italic>Pleosporales</italic> (<xref ref-type="bibr" rid="ref81">Mugambi &
Huhndorf 2009</xref>).</p></list-item><list-item><p>6. Ascospores (9–)10–12(–13) x 4–5(–6) μm;
cosmopolitan.......................................................................................................
<italic><bold>P. pusillum</bold></italic>6. Ascospores slightly
larger.........................................................................................................................................................................
7</p></list-item><list-item><p>7. Ascospores (10–)12–14(–18) x
(4–)5–7(–8) μm; ascomata +/- confluent laterally, in
parallel rows, semi-immersed to erumpent;
cosmopolitan................................................................................................................................
<italic><bold>P. lineare</bold></italic>7. Ascospores similar in length; ascomata not
confluent laterally, usually entirely
superficial......................................................................
8</p></list-item><list-item><p>8. Ascospores (10–)14–16(–18) x (4.5–)5–6
μm;
cosmopolitan.................................................................................................
<italic><bold>P. simulans</bold></italic>8. Ascospores slightly
larger.........................................................................................................................................................................
9</p></list-item><list-item><p>9. Ascospores (15–)16–18(–20) x 5–6(–7) um;
<italic>Sporidesmium stygium</italic> anamorph usually present; North and South
America,
Africa........................................................................................................................................
<italic><bold>P. clavisporum</bold></italic>9. Ascospores slightly larger in length and
breadth....................................................................................................................................
10</p></list-item><list-item><p>10. Ascospores (14–)16–18(–21) x
(6–)8–9(–10) μm; Europe, North
Africa..........................................................................
<italic><bold>P. chambianum</bold></italic>10. Ascospores slightly
larger.......................................................................................................................................................................
10</p></list-item><list-item><p>11. Ascospores 18–24 x 10–12 μm;
Argentina.......................................................................................................................
<italic><bold>P. uspallatense</bold></italic>11. Ascospores slightly
larger.......................................................................................................................................................................
12</p></list-item><list-item><p>12. Ascospores 25–32 x 5–8 μm, with a gelatinous sheath;
Japan................................................................................................
<italic><bold>P. sasicola</bold></italic>12. Ascospores slightly
larger.......................................................................................................................................................................
13</p></list-item><list-item><p>13. Ascospores 26–35 x 8–15 μm;
Chile......................................................................................................................................
<italic><bold>P. ephedrae</bold></italic>13. Ascospores
(59–)62–68(–76) x 13–15 μm; North and South
America......................................................................................
<italic><bold>O. curtisii</bold>Note</italic>: The genus <italic>Ostreichnion</italic>,
previously placed in the <italic>Mytilinidiaceae</italic>, has been transferred to the
<italic>Hysteriaceae</italic> (<xref ref-type="bibr" rid="ref18">Boehm <italic>et
al.</italic> 2009</xref>).</p></list-item><list-item><p>14. Hysterothecia usually borne in/on subicula, typically bifurcated,
forming radiating flabelliform or pseudo-stellate composites, with or without
a stroma (Type
III)...........................................................................................................................................
<italic><bold>Gloniaceae</bold>Note</italic>: In this study, a key to the species of the
<italic>Gloniaceae</italic> is provided under that family.
14. Hysterothecia not
bifurcated, forming radiating flabelliform or pseudo-stellate composites, nor
with a stroma...................................... 15</p></list-item><list-item><p>15. Ascospores less than 30 μm
long...........................................................................................................................................................
16
15. Ascospores more than 30 μm
long.........................................................................................................................................................
17</p></list-item><list-item><p>16. Ascospores (19–)22–25(–27) x
(6–)7–9(–10) μm, both ends acuminate, with a prominent
septal constriction; cosmopolitan (Type
II)..........................................................................................................................................................
<italic><bold>P. araucanum</bold></italic>16. Ascospores 22–28 x 4–6 μm,
acuminate, 1-septate, hyaline and with a mucilaginous sheath when young, but
acquiring additional septa and pigmentation with age, to become
3–5-septate and pale brown at maturity;
Kenya..................................................................................................................................................................................
<italic><bold>A. latirostrum</bold>Note</italic>: <italic>A. latirostrum</italic> lies within
the <italic>Pleosporales</italic> (<xref ref-type="bibr" rid="ref81">Mugambi &
Huhndorf 2009</xref>).</p></list-item><list-item><p>17. Ascospores 30–43 x 4–9.8 μm; Argentina (Type
II).................................................................................................................
<italic><bold>P. colihuae</bold></italic>17. Ascospores about 45 x 9 μm; Brazil (Type
II)......................................................................................................................
<italic><bold>P. hysterinum</bold></italic></p></list-item></list></p><p>Lee & Crous (<xref ref-type="bibr" rid="ref49">2003</xref>)
identified a series of isolates from South Africa on the <italic>Restionaceae</italic>as <italic>Glonium compactum</italic>(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=112412&link_type=cbs">CBS 112412</ext-link>, CMW
18760, CMW 17941). However, in their study they did not note the presence of
subicula, nor a stromal crust. These features were stressed for this taxon by
Zogg (<xref ref-type="bibr" rid="ref122">1962</xref>). These same
isolates were used in Boehm <italic>et al.</italic>(<xref ref-type="bibr" rid="ref18">2009</xref>), and were shown to
associate, with high branch support, with two species of <italic>Psiloglonium, P.
clavisporum</italic> and <italic>P. simulans</italic>, distant from the other species of
<italic>Glonium</italic> surveyed (<italic>e.g., G. stellatum</italic> and <italic>G.
circumserpens</italic>). Thus, a new combination was proposed, <italic>Psiloglonium
compactum</italic>. However, it is now realised that this new combination was made
in error and is hereby retracted. It must be concluded that the South African
isolates (<xref ref-type="bibr" rid="ref49">Lee & Crous
2003</xref>) were not <italic>G. compactum</italic>, due to the absence of
subicula and stroma, but rather, we suspect, the cosmopolitan <italic>P.
araucanum</italic>, which has similar, but slightly smaller, fusiform acuminate
didymospores. Lee & Crous
(<xref ref-type="bibr" rid="ref49">2003</xref>) give the ascospore
measurements for the South African “<italic>G. compactum</italic>” as
(24–)26–27(–30) x (4–)5–6(–7) μm, which
matches closely those given above for <italic>P. araucanum</italic>. Furthermore, the
illustrations in Lee & Crous
(<xref ref-type="bibr" rid="ref49">2003</xref>) closely match <italic>P.
araucanum</italic>, and not those of <italic>G. compactum</italic>, as given by Zogg
(<xref ref-type="bibr" rid="ref122">1962</xref>). If we are correct in
assuming that the South African isolates used in Boehm <italic>et al.</italic>(<xref ref-type="bibr" rid="ref18">2009</xref>) are in fact <italic>P.
araucanum</italic> (Type II) and not <italic>G. compactum</italic> (Type III), then this
would provide a high degree of support for the inclusion of species with
acuminate spore apices, belonging to Type II, in the genus
<italic>Psiloglonium</italic>, along with species with obtuse spore apices, belonging
to Type I (<italic>e.g., P. simulans</italic> and <italic>P. clavisporum</italic>). A
reanalysis of the original South African herbarium specimens from which the
sequences were derived (PREM 57570, PREM 57569, PREM 57566), by S.
Marincowitz, has confirmed that they do indeed correspond to <italic>P.
araucanum</italic> and not to <italic>G. compactum</italic>. Molecular data thus supports
the association of Types I and II within the genus <italic>Psiloglonium</italic>.</p><p>In addition to the 12 currently recognised species in
<italic>Psiloglonium</italic>, the following key also includes entries for the
unrelated <italic>Gloniaceae, Anteaglonium</italic> and <italic>Ostreichnion
curtisii</italic>.</p><p><italic><bold>Actidiographium</bold></italic> Lar.N. Vassiljeva<bold>,</bold> Mikol. Fitopatol.
34 (6): 4. 2000.</p><p>Vasilyeva (<xref ref-type="bibr" rid="ref116">2000</xref>)
established the monotypic genus <italic>Actidiographium</italic> to accommodate a
hysteriaceous fungus with pigmented one-septate ascospores, reminiscent of
those found in <italic>Actidium</italic> in the <italic>Mytilinidiaceae</italic>. However, in
<italic>Actidiographium orientale</italic>, the two-celled spores are borne in a
typical thick-walled hysterothecium. The pigmented didymospores measure
13.2–16.5 x 3–4 μm. Molecular data are lacking for this
taxon.</p><p><italic><bold>Hysterocarina</bold></italic> H. Zogg, Ber. Schweiz. Bot. Ges. 59: 39.
1949.</p><p>Zogg (<xref ref-type="bibr" rid="ref121">1949</xref>) erected this
monotypic genus for <italic>Hysterocarina paulistae</italic>, with pigmented
dictyospores as in <italic>Hysterographium</italic>, but the hysterothecia are borne
within the substrate, barely erumpent at maturity, and with a cristate,
slightly evaginated longitudinal keel, instead of the invaginated sulcus
typical of most members of the <italic>Hysteriaceae</italic>. Described from old wood
of <italic>Eucalyptus</italic> sp. in Brazil, the pigmented dictyospores measure
20–25 x 8–10 μm. The presence of an evaginated keel-like
fissure in <italic>Hysterocarina</italic> is intriguing, as it seems to belong to an
evolutionary trend that culminates in the <italic>Mytilinidiaceae</italic> and
<italic>Gloniaceae</italic>. Clearly, molecular data are needed to resolve these
issues.</p><p><italic><bold>Ostreichnion</bold></italic> Duby, Mém. Soc. Phys. Genève 16:
22. 1862.</p><p><list list-type="simple"><list-item><p>= <italic>Ostreion</italic> Sacc., Syll. Fung. 2: 765. 1883.</p></list-item></list></p><p>Since its reappraisal (<xref ref-type="bibr" rid="ref4">Barr
1975</xref>), the genus <italic>Ostreichnion</italic> has been heterogeneous, due
to the inclusion of <italic>O. curtisii</italic> an unusual taxon, from the
southeastern United States (<xref ref-type="bibr" rid="ref61">Lohman
1937</xref>) and Brazil (<xref ref-type="bibr" rid="ref122">Zogg
1962</xref>). It is very different from the other two species of this
genus, namely the type <italic>O. sassafras</italic> and <italic>O. nova-caesariense</italic>.
Whereas the latter two species possess pigmented dictyospores, in <italic>O.
curtisii</italic> the ascospores are 1-septate below the middle, with walls
greatly thickened towards the spore apices. When mounted under different
stains, the spore cytoplasm appears subdivided into numerous compartments,
giving the impression of a potentially muriform structure. Lohman
(<xref ref-type="bibr" rid="ref61">1937</xref>) provided details as to
the highly unusual spore germination process in this fungus, which involves a
distended apical plug and numerous median germ tubes, differing from that
found in species of <italic>Psiloglonium</italic> and <italic>Glonium</italic>, which send out
apical germ tubes (Lohman
<xref ref-type="bibr" rid="ref55">1931</xref>,
<xref ref-type="bibr" rid="ref56">1932a</xref>). <italic>Ostreichnion
sassafras</italic> occurs on both sides of the Atlantic, as well as in China, and
has been recovered from <italic>Sassafras, Quercus, Liriodendron</italic>, and
<italic>Liquidambar</italic> (<xref ref-type="bibr" rid="ref15">Bisby
1932</xref>, <xref ref-type="bibr" rid="ref111">Teng 1933</xref>,
<xref ref-type="bibr" rid="ref4">Barr 1975</xref>). It is unusual in
having very large dictyospores, measuring (65–)76–100(–135)
x 20–32 μm, with up to 27 septa, borne four to an ascus.
<italic>Ostreichnion nova-caesariense</italic> is known only from the type locality in
New Jersey on <italic>Pinus</italic>, and has similar, but smaller, ascospores
(<xref ref-type="bibr" rid="ref4">Barr 1975</xref>).</p></sec><sec><title>Key to the species of <italic>Ostreichnion</italic></title><p><list list-type="simple"><list-item><p>1. Ascospores mostly 1-septate, ends greatly thickened,
(45–)62–80 x (10–)12–15 μm; North & South
America........................ <italic><bold>O. curtisii</bold></italic>1. Ascospores
with both transverse and longitudinal
septa..........................................................................................................................
2</p></list-item><list-item><p>2. Ascospores measuring 35–45(–50) x 11–13 μm, with
7–13 septa, borne eight to an ascus; North America.......... <italic><bold>O.
nova-caesariense</bold></italic>2. Ascospores measuring
(65–)76–100(–135) x 20–32 μm, with up to 27 septa,
borne four to an ascus; cosmopolitan......... <italic><bold>O. sassafras</bold></italic></p></list-item></list></p><p>Based on a recent four-gene analysis
(<xref ref-type="bibr" rid="ref18">Boehm <italic>et al.</italic> 2009</xref>),
the genus <italic>Ostreichnion</italic>, previously in the <italic>Mytilinidiaceae</italic>(Barr <xref ref-type="bibr" rid="ref4">1975</xref>,
<xref ref-type="bibr" rid="ref8">1990a</xref>), was transferred to the
<italic>Hysteriaceae</italic>. This was based on sequence data derived from two of the
three species (<xref ref-type="table" rid="tbl1">Table 1</xref>), namely
<italic>O. curtisii</italic> (<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=198.34&link_type=cbs">CBS
198.34</ext-link>) and <italic>O. sassafras</italic>(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=322.34&link_type=cbs">CBS 322.34</ext-link>),
deposited by Lohman in 1934. Although both species find residency within Clade
C (<xref ref-type="fig" rid="fig1">Fig. 1</xref>), their association
with the genus <italic>Hysterium</italic> could not have been predicted. Given the
unique nature of the ascospore in <italic>O. curtisii</italic>, considered potentially
muriform, one would assume affinities with the genus <italic>Hysterographium
sensu</italic> Zogg (<xref ref-type="bibr" rid="ref122">1962</xref>), or,
given its 1-septate ascospores at maturity, with <italic>Psiloglonium</italic>, where
it was originally treated by Lohman
(<xref ref-type="bibr" rid="ref61">1937</xref>) as <italic>Glonium
curtisii</italic>. However, molecular data suggest neither. Instead, <italic>O.
curtisii</italic> shares a subclade with <italic>Hysterium barrianum</italic>, with
9-septate phragmospores (<xref ref-type="fig" rid="fig1">Fig.
1</xref>). <italic>Ostreichnion sassafras</italic> is more distant within Clade
C. Although we recognise the genus as artificial, we present the following
key, adapted from Barr (<xref ref-type="bibr" rid="ref4">1975</xref>),
to facilitate species identification.</p><p><italic><bold>Rhytidhysteron</bold></italic> Speg., Anales Soc. Ci. Argent. 12: 188.
1881.</p><p>The genus <italic>Rhytidhysteron</italic> is characterised by ascomata that are at
first closed and navicular (<italic>e.g.</italic>,
<xref ref-type="fig" rid="fig10">Fig. 10K</xref>), somewhat resembling
those found in the <italic>Hysteriaceae</italic>, but then later opening by a
longitudinal sulcus to become irregularly apothecioid at maturity, often with
incurved margins (<italic>e.g.</italic>, <xref ref-type="fig" rid="fig10">Fig.
10M</xref>) – a feature never observed in the
<italic>Hysteriaceae</italic>. The peridium in <italic>Rhytidhysteron</italic> is somewhat
gelatinous when wet, as compared to the hard, carbonaceous peridium found in
the <italic>Hysteriaceae</italic>. Although ascomata may possess striations, in
<italic>Rhytidhysteron</italic> these are perpendicular to the long axis
(<xref ref-type="fig" rid="fig10">Fig. 10K</xref>), rather than
parallel, as in the <italic>Hysteriaceae</italic> (<italic>e.g.</italic>, Figs
<xref ref-type="fig" rid="fig1">1A</xref>,
<xref ref-type="fig" rid="fig2">2B</xref>, and
<xref ref-type="fig" rid="fig6">6A</xref>). The ascospores in
<italic>Rhytidhysteron</italic> tend to be heavily pigmented and thick-walled, as
opposed to lightly pigmented and thin-walled in the <italic>Hysteriaceae.</italic>These features, among others, have been used to place <italic>Rhytidhysteron</italic>within the <italic>Patellariaceae</italic> (<italic>e.g.</italic>,
<xref ref-type="bibr" rid="ref48">Kutorga & Hawksworth
1997</xref>). Samuels & Müller
(<xref ref-type="bibr" rid="ref97">1979</xref>) revised the genus,
providing a number of synonyms, and accepted only two species, namely the
type, <italic>R. rufulum</italic> (<xref ref-type="fig" rid="fig10">Fig.
10E–K</xref>), with 3-septate phragmospores, and <italic>R.
hysterinum</italic> (<xref ref-type="fig" rid="fig10">Fig. 10M</xref>),
with 1-septate spores, both darkly pigmented and thick-walled. Anamorphs have
been characterised as <italic>Diplodia</italic>- and <italic>Aposphaeria</italic>-like
(<xref ref-type="bibr" rid="ref97">Samuels & Müller
1979</xref>). Subsequently, another two species have been accepted in the
genus, namely <italic>R. dissimile</italic>(<xref ref-type="bibr" rid="ref72">Magnes 1997</xref>), with 5-septate
phragmospores, and <italic>R. opuntiae</italic> (1990b), from the American South West,
with short pigmented dictyospores (<xref ref-type="fig" rid="fig10">Fig.
10A–D</xref>), reminiscent of those found in <italic>Hb. mori</italic>.</p><p>Dictyospores of both <italic>R. opuntiae</italic> and <italic>Hb. mori</italic> are similar
in shape, obovoid, with obtuse ends, and are also similar in size and
septation. In both, the longitudinal septum is usually associated with the
mid-cells, but on occasion it can be found obliquely in the end cells.
However, unlike <italic>Hb. mori</italic>, the spores of <italic>R. opuntiae</italic> are
thick-walled, verruculose and darkly pigmented. The most surprising
morphological feature of <italic>R. opuntiae</italic> is that the spores are not borne
within patellarioid ascomata, as in other members of the genus. Rather, the
ascomata are hysterithecioid, that is, carbonaceous, navicular, with an
invaginated longitudinal sulcus (<xref ref-type="fig" rid="fig10">Fig.
10A–B</xref>). In hindsight, it is remarkable that Barr (1990)
recognised <italic>R. opuntiae</italic> as a member of <italic>Rhytidhysteron</italic>,
transferring it from <italic>Hysterographium opuntiae,</italic> despite the presence
of hysterithecioid ascomata. In this study we were fortunate to acquire an
isolate of <italic>R. opuntiae</italic> from Kenya (GKM 1190 / BPI 879805).
<italic>Rhytidhysteron opuntiae</italic> falls distant from <italic>R. rufulum</italic> and
<italic>R. hysterinum,</italic> lying outside of Clade E altogether
(<xref ref-type="fig" rid="fig1">Fig. 1</xref>). Although both
morphological and molecular data suggest that <italic>R. opuntiae</italic> should be
removed from the genus <italic>Rhytidhysteron</italic>, this is based only on a single
specimen, and clearly needs to be substantiated with other isolates.</p><p>The six isolates of <italic>R. rufulum</italic> included one from Kenya (GKM 361A /
BPI 879806; <xref ref-type="fig" rid="fig10">Fig. 10E–J</xref>),
four from Ghana (EB 0381 / BPI 879807,
<xref ref-type="fig" rid="fig10">Fig. 10L</xref>; EB 0382 / BPI
879808, <xref ref-type="fig" rid="fig10">Fig. 10K</xref>; EB 0383 /
879809; EB 0384 / BPI 879810), and one from Europe
(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=306.38&link_type=cbs">CBS 306.38</ext-link>). Also
included was one isolate of <italic>R. hysterinum</italic> from France (EB 0351 / BPI
879804). Three of the Ghanian isolates clustered together in Clade E
(<xref ref-type="fig" rid="fig1">Fig. 1</xref>), but one (EB 0381 /
BPI 879807) associated in another subclade, along with the Kenyan (GKM 361A)
and European (<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=306.38&link_type=cbs">CBS
306.38</ext-link>) accessions of <italic>R. rufulum</italic>. The morphology of the
ascomata (<xref ref-type="fig" rid="fig10">Fig. 10L</xref>) of <italic>R.
rufulum</italic> EB 0381 (BPI 879807) differs from other more typical specimens of
<italic>R. rufulum</italic> (<italic>e.g.</italic>, <xref ref-type="fig" rid="fig10">Fig.
10 K</xref>), although the 3-septate spores in both are identical.
Finally, molecular data indicate that <italic>R. hysterinum</italic>, with 1-septate
spores, falls outside of the <italic>R. rufulum</italic> subclades, while still within
Clade E (<xref ref-type="fig" rid="fig1">Fig. 1</xref>).</p><p><fig position="float" id="fig10"><label>Fig. 10.</label><caption><p>The genus <italic>Rhytidhysteron</italic> (Clade E). A–D. <italic>Rhytidhysteron
opuntiae</italic> [GKM 1190 (BPI 879805), Kenya]; E–J. <italic>Rhytidhysteron
rufulum</italic> [GKM 361A (BPI 879806), Kenya]; K. <italic>Rhytidhysteron
rufulum</italic> [EB 0382 (BPI 879808), Ghana]; L. <italic>Rhytidhysteron rufulum</italic>[EB 0381 (BPI 879807), Ghana]; M. <italic>Rhytidhysteron hysterinum</italic> [EB 0351
(BPI 879804) France, photo by Alain Gardiennet]. Scale bar (habitat) = 1 mm;
Scale bar (spores and asci) = 10 μm.</p></caption><graphic xlink:href="49fig10"></graphic></fig></p><p>Boehm <italic>et al.</italic>(<xref ref-type="bibr" rid="ref18">2009</xref>) were the first to
provide sequence data indicating that <italic>Rhytidhysteron</italic> does not lie
within the <italic>Patellariaceae</italic>. Although initially based on only a single
isolate of <italic>R. rufulum</italic>(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=306.38&link_type=cbs">CBS 306.38</ext-link>), the
genus was tentatively noted to be associated with the <italic>Hysteriaceae</italic>.
In the current study, a total of eight isolates, representing three species,
clearly indicates that that the genus <italic>Rhytidhysteron</italic> belongs to the
family <italic>Hysteriaceae</italic>, and not to the <italic>Patellariaceae</italic>, the
latter defined in this study to include <italic>Hysteropatella clavispora</italic>(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=247.34&link_type=cbs">CBS 247.34</ext-link>),
<italic>Hp. elliptica</italic> (<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=935.97&link_type=cbs">CBS
935.97</ext-link>), and <italic>Patellaria atrata</italic>(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=958.97&link_type=cbs">CBS 958.97</ext-link>).</p><p>Earlier, Barr (<xref ref-type="bibr" rid="ref7">1987</xref>) had
noted the differences between <italic>Rhytidhysteron</italic> and other members of the
<italic>Patellariaceae</italic>, stating: “<italic>Rhytidhysteron rufulum</italic>illustrates the problem: paraphysoids and a well-developed pseudoepithecium
are conspicuous, but the structure of the peridium, thickened base of ascoma,
cylindric asci, are all features attributed to members of the
<italic>Hysteriaceae</italic>. When the heterogeneous family <italic>Patellariaceae</italic>is revised, <italic>Rhytidhysteron</italic> should be segregated in its own
family”. Samuels & Müller
(<xref ref-type="bibr" rid="ref97">1979</xref>) also noted that
“The genus does not have any close relatives in the heterogeneous
<italic>Patellariaceae</italic>”. However, other authors
(<xref ref-type="bibr" rid="ref13">Bezerra & Kimbrough 1982</xref>)
presented arguments against the inclusion of <italic>Rhytidhysteron</italic> within
the <italic>Hysteriaceae</italic>, based on patterns of centrum development.
Nevertheless, molecular data presented here, necessitate a radical reappraisal
of the <italic>Hysteriaceae</italic> to include patellarioid forms.</p></sec><sec><title>Key to the species of <italic>Rhytidhysteron</italic></title><p><list list-type="simple"><list-item><p>1. Ascospores mainly 1-septate;
Europe................................................................................................................................
<italic><bold>R. hysterinum</bold></italic>1. Ascospores with more than one
septum...................................................................................................................................................
2</p></list-item><list-item><p>2. Ascospores mainly
3-septate....................................................................................................................................................................
3
2. Ascospores with five or more septa;
Europe..........................................................................................................................
<italic><bold>R. dissimile</bold></italic></p></list-item><list-item><p>3. Ascospores with three transverse, but also one or more longitudinal
septa; Southwestern United States, East Africa......... <italic><bold>R.
opuntiae</bold></italic>3. Ascospores transversely 3-septate, with no
longitudinal septa;
cosmopolitan.........................................................................
<italic><bold>R. rufulum</bold></italic></p></list-item></list></p><p><italic><bold>Mytilinidiaceae</bold></italic> Kirschst. 1924,
<italic><bold>Mytilinidiales</bold></italic> E.W.A. Boehm, C.L. Schoch & J.W. Spatafora
2009.</p><p><list list-type="simple"><list-item><p>= <italic>Lophiaceae</italic> H. Zogg <italic>ex</italic> Arx & E. Müll., Stud.
Mycol. 9: 60. 1975.</p></list-item><list-item><p>≡ <italic>Lophiaceae</italic> H. Zogg, Beitr. Kryptogamenfl. Schweiz. 11(3):
90. 1962, <italic>nom. inval.</italic> ICBN Art. 36.</p></list-item></list></p><p>Fungi classified in the <italic>Mytilinidiaceae</italic>(<xref ref-type="bibr" rid="ref46">Kirschstein 1924</xref>) are
characterised by fragile yet persistent carbonaceous ascomata, which range
from globoid to obovoid to strongly laterally compressed erect, bivalve
shell-shaped (<italic>i.e.</italic>, conchate) structures, standing on edge, with
lateral walls more or less connivent, and extended vertically, in some
species, to a prominent longitudinal keel or cristate apex. Mytilinidioid
fungi possess a thin-walled, scleroparenchymatous peridium enclosing a
hamathecium of narrow trabeculate pseudoparaphyses, borne in a gel matrix,
which are often sparse to lacking at maturity. Bitunicate asci are borne in a
basal, rarely lateral orientation within the centrum, and contain eight,
rarely four, ascospores, overlapping uniseriate, biseriate or in one or two
fascicles. Ascospores are diverse, ranging from scolecospores to didymospores
to phragmospores or dictyospores, hyaline, soon yellow to dark brown, and
generally showing bipolar symmetry (<xref ref-type="bibr" rid="ref122">Zogg
1962</xref>, Barr <xref ref-type="bibr" rid="ref7">1987</xref>,
<xref ref-type="bibr" rid="ref8">1990a</xref>). Anamorphs in the
<italic>Mytilinidiaceae</italic> are primarily coelomycetous (<italic>e.g., Aposphaeria,
Pyrenochaeta, Camaroglobulus, Dothiorella</italic>-like, and
<italic>Sclerochaeta</italic>) and less frequently hyphomycetous (<italic>e.g.,
Chalara</italic>-like, <italic>Papulaspora</italic>, and <italic>Septonema</italic>) (Lohman
<xref ref-type="bibr" rid="ref57">1932b</xref>,
<xref ref-type="bibr" rid="ref58">1933a</xref>,
<xref ref-type="bibr" rid="ref59">b</xref>,
<xref ref-type="bibr" rid="ref17">Blackwell & Gilbertson
1985</xref>, <xref ref-type="bibr" rid="ref106">Speer
1986</xref>). Typically temperate in distribution, mytilinidioid fungi
are found in association with the wood, bark, resin, cones, scales, needles,
seeds, and roots of gymnosperms.</p><p>Currently accepted genera in the <italic>Mytilinidiaceae</italic> include:
<italic>Actidium, Lophium, Mytilinidion, Ostreola</italic>, and <italic>Quasiconcha</italic>,
to which has recently been added <italic>Zoggium</italic>(<xref ref-type="bibr" rid="ref57">Lohman 1932b</xref>,
<xref ref-type="bibr" rid="ref122">Zogg 1962</xref>,
<xref ref-type="bibr" rid="ref26">Darker 1963</xref>, Barr
<xref ref-type="bibr" rid="ref4">1975</xref>,
<xref ref-type="bibr" rid="ref8">1990a</xref>,
<xref ref-type="bibr" rid="ref11">Barr & Blackwell 1980</xref>,
<xref ref-type="bibr" rid="ref117">Vasilyeva 2001</xref>). The genus
<italic>Ostreichnion</italic>, previously classified within the
<italic>Mytilinidiaceae</italic>, has been removed to the <italic>Hysteriaceae</italic>(<xref ref-type="bibr" rid="ref18">Boehm <italic>et al.</italic> 2009</xref>).
The genus <italic>Glyphium</italic>, originally classified within the
<italic>Mytilinidiaceae</italic>, has recently been transferred to the
<italic>Chaetothyriales</italic> in the <italic>Eurotiomycetes</italic>(<xref ref-type="bibr" rid="ref52">Lindemuth <italic>et al.</italic>2001</xref>, <xref ref-type="bibr" rid="ref66">Lumbsch <italic>et al.</italic>2005</xref>). This has been restated in a number of subsequent
publications (<xref ref-type="bibr" rid="ref63">Lücking <italic>et al.</italic>2004</xref>, <xref ref-type="bibr" rid="ref98">Schmitt <italic>et al.</italic>2005</xref>, <xref ref-type="bibr" rid="ref37">Geiser <italic>et al.</italic>2006</xref>, <xref ref-type="bibr" rid="ref47">Kodsueb <italic>et al.</italic>2006</xref>), including the Assembling the Fungal Tree of Life (AFTOL)
Project (<xref ref-type="bibr" rid="ref71">Lutzoni <italic>et al.</italic>2004</xref>). A study currently in preparation (Boehm <italic>et al</italic>.)
will address issues related to the phylogenetic placement of the genus
<italic>Glyphium</italic>. Despite their transference out of the
<italic>Mytilinidiaceae</italic>, both <italic>Ostreichnion</italic> and <italic>Glyphium</italic> are
included in the current key to effectuate identification of morphologically
similar fungi, regardless of whether close phylogeny is implied or not.</p></sec><sec><title>Key to the genera of the <italic>Mytilinidiaceae</italic></title><p><list list-type="simple"><list-item><p>1. Ascospores 1-septate, small, shorter than 30
μm.....................................................................................................................................
2
1. Ascospores not didymospores, or if 1-septate, then longer than 30
μm...................................................................................................
3</p></list-item><list-item><p>2. Didymospores brown, ellipsoid, symmetric, with coarsely reticulate wall;
6–8 x 5–5.5 μm.................................................
<italic><bold>Quasiconcha</bold></italic>2. Didymospores olive- to reddish brown, walls
thin, smooth or delicately longitudinally striate, but not reticulated; longer
than 10
μm........................................................................................................................................
<italic><bold>Actidium</bold></italic></p></list-item><list-item><p>3. Ascospores filiform, multi-septate, about equal in length to the ascus,
in some case, at maturity longer than the ascus, often spirally
arranged..............................................................................................................................................................................
4
3. Ascospores ellipsoid, fusoid, cylindrical, if scolecospores, then
shorter than the ascus and not spirally
arranged.................................. 6</p></list-item><list-item><p>4. Ascomata conchate, solitary to gregarious, but never forming fused,
ridge-like assemblages...................................................
<italic><bold>Lophium</bold></italic>4. Ascomata either forming rigid, fused band- or
ridge-like structures or solitary, erect, dolabrate to
ligulate.............................................. 5</p></list-item><list-item><p>5. Ascomata densely gregarious, forming band- or ridge-like
assemblages...................................................................................
<italic><bold>Zoggium</bold></italic>5. Ascomata erect, dolabrate to ligulate in
outline; often with subtending hyphal strands;
cosmopolitan......................................
<italic><bold>Glyphium</bold>Note</italic>: A key to the species is not presented
here.</p></list-item><list-item><p>6. Ascospores transversely septate phragmospores, or
scolecospores.....................................................................................
<italic><bold>Mytilinidion</bold></italic>6. Ascospores dictyospores, or large and
remaining
1-septate....................................................................................................................
7</p></list-item><list-item><p>7. Ascospores ellipsoid, less than 30 μm long, with a single
longitudinal septum, usually passing through the mid-cells, or spanning the
entire length of the
ascospore............................................................................................................................
<italic><bold>Ostreola</bold></italic>7. Ascospores ellipsoid or cylindric, longer than
30 μm, with several longitudinal septa in cells or large and remaining
1-septate.......................................................................................................................................................
<italic><bold>Ostreichnion</bold>Note</italic>: The genus <italic>Ostreichnion</italic>previously classified within the <italic>Mytilinidiaceae</italic>(<xref ref-type="bibr" rid="ref8">Barr 1990a</xref>) has been
transferred to the <italic>Hysteriaceae</italic>(<xref ref-type="bibr" rid="ref18">Boehm <italic>et al.</italic>2009</xref>).</p></list-item></list></p><p><italic><bold>Actidium</bold></italic> Fr., Observ. Mycol. 1: 190. 1815.</p><p><list list-type="simple"><list-item><p>= <italic>Mytilinidion</italic> subgen. <italic>Bulliardella</italic> Sacc., Syll. Fung. 2:
764. 1883.</p></list-item><list-item><p>= <italic>Bulliardella</italic> (Sacc.) Paoli, Nuovo Giorn. Bot. Ital. 12:101.
1905.</p></list-item><list-item><p>= <italic>Ostreionella</italic> Seaver, Sci. Surv. Porto Rico & Virgin Islands
8(1): 77. 1926.</p></list-item></list></p><p>The genus <italic>Actidium</italic> was established by Fries
(<xref ref-type="bibr" rid="ref34">1823</xref>) to accommodate <italic>A.
hysterioides</italic>, a stellate mytilinidioid fungus found on <italic>Pinus</italic> and
<italic>Picea</italic> in Europe, with two-celled, symmetric ascospores, light olive-
to reddish-brown, later noted to be faintly longitudinally striate
(<xref ref-type="bibr" rid="ref8">Barr 1990a</xref>). Fries
(<xref ref-type="bibr" rid="ref34">1823</xref>) noted its similarity
with the genus <italic>Glonium</italic>. Zogg
(<xref ref-type="bibr" rid="ref122">1962</xref>) recognised a total of
four species, namely <italic>A. hysterioides, A. baccarinii</italic>, both from
Europe, <italic>A. pulchra</italic>, from China, and <italic>A. nitidum</italic>, from Europe
and North America, on <italic>Pinus, Picea, Juniperus</italic>, and <italic>Thuja</italic>(<xref ref-type="bibr" rid="ref122">Zogg 1962</xref>,
<xref ref-type="bibr" rid="ref8">Barr 1990a</xref>). Due to
similarities in ascospore morphology, the genus <italic>Actidium</italic> may have
affinities with other didymospored hysteriaceous genera (<italic>e.g.,
Actidiographium, Glonium</italic> and <italic>Psiloglonium</italic>), although molecular
data are presently lacking.</p></sec><sec><title>Key to the species of <italic>Actidium</italic></title><p><list list-type="simple"><list-item><p>1. Ascomata stellate; spores 11–14 x (1.5–)2–3 μm; on
<italic>Pinus, Picea</italic>,
Europe....................................................................
<italic><bold>A. hysterioides</bold></italic>1. Ascomata shell-shaped (conchate), not
star-shaped................................................................................................................................
2</p></list-item><list-item><p>2. Ascospores (9–)11–14(–16) x (1.5–)2–3
μm; on <italic>Pinus, Picea, Juniperus</italic>, Europe, North
America......................................... <italic><bold>A.
nitidum</bold></italic>2. Ascospores
larger.....................................................................................................................................................................................
3</p></list-item><list-item><p>3. Ascospores (16–)18–22(–24) x
(3–)4–5(–6) μm; on <italic>Pinus, Picea, Thuja,</italic>Europe...............................................................
<italic><bold>A. baccarinii</bold></italic>3. Ascospores 23–28 x 6–7.5
μm;
China......................................................................................................................................
<italic><bold>A. pulchra</bold></italic></p></list-item></list></p><p><italic><bold>Quasiconcha</bold></italic> M.E. Barr & M. Blackw., Mycologia 72: 1224.
1980.</p><p>The genus <italic>Quasiconcha</italic> was established by Barr & Blackwell
(<xref ref-type="bibr" rid="ref11">1980</xref>) to accommodate <italic>Q.
reticulata</italic>, an unusual mytilinidioid fungus, with 1-septate, highly
reticulated ascospores, borne in conchate, thin-walled ascomata, found in
association with <italic>Juniperus</italic> seeds excreted in dung and the roots of
two conifers from the southwestern United States
(<xref ref-type="bibr" rid="ref11">Barr & Blackwell 1980</xref>,
<xref ref-type="bibr" rid="ref17">Blackwell & Gilbertson
1985</xref>). In the present study, we were fortunate to obtain original
material (RLG 141189) of <italic>Q. reticulata</italic>(<xref ref-type="table" rid="tbl1">Table 1</xref>) from Meredith
Blackwell (Louisiana State University, Baton Rouge, LA), from which we
isolated DNA (EB QR). Sequence data (<xref ref-type="fig" rid="fig1">Fig.
1</xref>) clearly indicate that the genus <italic>Quasiconcha</italic> belongs to
the <italic>Mytilinidiaceae</italic>, in close association with <italic>Lophium</italic>, to
which its fruitbodies most closely resemble.</p><p><italic><bold>Mytilinidion</bold></italic> Duby, Mém. Soc. Phys. Genève 16:
34. 1862.</p><p><list list-type="simple"><list-item><p>= <italic>Mytilidion</italic> Sacc., Atti Soc. Veneto-Trentino Sci. Nat. Padova 4:
99. 1875.</p></list-item><list-item><p>= <italic>Hypodermopsis</italic> Earle, Bull. New York Bot. Gard. 2: 345. 1902.</p></list-item><list-item><p>= <italic>Murashkinskija</italic> Petr., Hedwigia 68: 203. 1928.</p></list-item></list></p><p>The genus <italic>Mytilinidion</italic>, the type for the family
<italic>Mytilinidiaceae</italic>, was established by Duby
(<xref ref-type="bibr" rid="ref31">1862</xref>) with an etymology from
<italic>Mytilus</italic>, a genus of mussels. Saccardo
(<xref ref-type="bibr" rid="ref96">1883</xref>, p. 760) considered the
name <italic>Mytilinidion</italic> to be linguistically incorrect and replaced it with
<italic>Mytilidion</italic>. It remained for Barr
(<xref ref-type="bibr" rid="ref4">1975</xref>) to note that the name
<italic>Mytilinidion</italic> had historical precedence
(<xref ref-type="bibr" rid="ref93">Rogers 1953</xref>), and therefore
should replace the later name <italic>Mytilidion</italic>. Species of
<italic>Mytilinidion</italic> are characterised by yellow- to reddish-brown,
ellipsoid, fusoid, obovoid to elongate, transversely septate, usually
symmetric, ascospores, or scolecospores, borne in thin-walled globoid to
conchate pseudothecia, with lateral walls more or less connivent and extended
vertically to a cristate apex. There are currently 15 recognised species,
occurring on the <italic>Pinaceae, Cupressaceae</italic>, and <italic>Taxodiaceae</italic>(<xref ref-type="bibr" rid="ref57">Lohman 1932b</xref>,
<xref ref-type="bibr" rid="ref122">Zogg 1962</xref>,
<xref ref-type="bibr" rid="ref106">Speer 1986</xref>,
<xref ref-type="bibr" rid="ref8">Barr 1990a</xref>).</p><p>Ascospore morphology can be used to discern four morphological types within
the genus, listed here by increasing ascospore length: (1) Short squat
phragmospores: <italic>M. acicola, M. resinae, M. decipiens, M. tortile</italic>(<xref ref-type="fig" rid="fig11">Fig. 11A–B</xref>), and <italic>M.
resinicola</italic>; (2) Elongate phragmospores, with a spore length to width
ratio of 10: 1 or less: <italic>M. californicum, M. mytilinellum</italic>(<xref ref-type="fig" rid="fig11">Fig. 11C–D</xref>), <italic>M.
rhenanum</italic>, and <italic>M. gemmigenum</italic>; (3) Fusoid or spindle-shaped
spores: <italic>M. thujarum, M. oblongisporum</italic>, and <italic>M. andinense</italic>; and
(4) Highly elongated phragmospores, termed scolecospores, with a length to
width ratio of 20: 1: <italic>M. scolecosporum, M. parvulum</italic> and <italic>M.
australe</italic> (<xref ref-type="fig" rid="fig11">Fig.
11E–I</xref>). These last three scolecosporous species were
postulated to form a transitional series to connect <italic>Mytilinidion</italic> with
the heretofore somewhat isolated genus <italic>Lophium</italic>(<xref ref-type="fig" rid="fig11">Fig. 11J–K</xref>), and formed
the basis for subgenus <italic>Lophiopsis</italic>, distinct from subgenus
<italic>Eu-Mytilinidion sensu</italic> Lohman
(<xref ref-type="bibr" rid="ref57">Lohman 1932b</xref>), a concept
accepted by Zogg (<xref ref-type="bibr" rid="ref122">1962</xref>).</p><p><fig position="float" id="fig11"><label>Fig. 11.</label><caption><p>The <italic>Mytilinidiaceae</italic>. A–B. <italic>Mytilinidion tortile</italic> [EB
0377 (BPI 879798), France]; C–D. <italic>Mytilinidion mytilinellum</italic> [EB
0386 (BPI 879796), France]; E–I. <italic>Mytilinidion australe</italic> [ANM
1524 (ILLS), U.S.A.; not incl.]; J–K. <italic>Lophium mytilinum</italic>[<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123344&link_type=cbs">CBS 123344</ext-link> (BPI
878736), U.S.A.]. Photo credits Alain Gardiennet, Figs. A–D. Scale bar
(habitat) = 500 μm; Scale bar (spores and asci) = 10 μm.</p></caption><graphic xlink:href="49fig11"></graphic></fig></p></sec><sec><title>Key to the species of <italic>Mytilinidion</italic></title><p><list list-type="simple"><list-item><p>1. Spore length to width ratio = 10: 1 or less (phragmospores): Subgenus
<italic>Eu-Mytilinidion sensu</italic> Lohman
(<xref ref-type="bibr" rid="ref57">1932b</xref>)...................................
2
1. Spore length to width ratio = approx. 20: 1 (scolecospores): Subgenus
<italic>Lophiopsis sensu</italic> Lohman
(<xref ref-type="bibr" rid="ref57">1932b</xref>).......................................
13</p></list-item><list-item><p>2. Ascomata not conchate, but erect, low and spreading at the base
(scutate), seated on a shield-like process fused to the substrate, apical
portion slightly connivent; ascospores
3–5(–6)-septate.......................................................................................
3
2. Ascomata conchate, standing on edge, usually with a clearly defined
longitudinal cristate
apex............................................................ 4</p></list-item><list-item><p>3. Ascospores 23–25 x 4–4.5(–5) μm, 3-septate;
California on
<italic>Sequoia</italic>.............................................................................
<italic><bold>M. californicum</bold></italic>3. Ascospores 14–22(–28) x
(4.5–)6–8(–10) μm, 3–4–5–(–6)
septate; on <italic>Juniperus, Thuja</italic>, Europe and North
America.....................................................................................................................................................................
<italic><bold>M. acicola</bold></italic></p></list-item><list-item><p>4. Ascospores elongate phragmospores, usually not constricted at the
septa.............................................................................................
5
4. Ascospores shorter, squat, or longer, but not narrowly elongated,
usually constricted at median
septum............................................... 7</p></list-item><list-item><p>5. Ascospores (2–)3(–5)-septate, measuring
(14–)16–22(–24) x (2.5–)3–4(–5) μm;
cosmopolitan..................................... <italic><bold>M.
mytilinellum</bold></italic>5. Ascospores longer, with more
septa.........................................................................................................................................................
6</p></list-item><list-item><p>6. Ascospores 3–5(–7)-septate, measuring
(24–)30–42(–50) x 3–5 μm;
Europe....................................................................
<italic><bold>M. rhenanum</bold></italic>6. Ascospores slightly curved, asymmetric,
(3–)7–9(–11)-septate, measuring
(27–)32–38(–48) x (4–)5–6(–8) μm;
cosmopolitan..................................................................................................................................................................
<italic><bold>M. gemmigenum</bold></italic></p></list-item><list-item><p>7. Ascospores (2–)3-septate, small, 10–13 x 4–6 μm;
resinicolous on <italic>Araucaria</italic>,
Brazil............................................................. <italic><bold>M.
resinae</bold></italic>7. Ascospores 3(–5)-septate,
longer.............................................................................................................................................................
8</p></list-item><list-item><p>8. Ascospores 3-septate, slightly curved, oblong-elliptic, with obtuse
ends, unconstricted, measuring (11–)13–15(–21) x
3–4(–6) μm; on <italic>Larix, Juniperus</italic>,
Europe.............................................................................
<italic><bold>M. decipiens</bold></italic>8. Ascospores longer, or similar in length but
then slightly
wider..................................................................................................................
9</p></list-item><list-item><p>9. Ascospores 3-septate, slightly curved, but oblong, fusiform, with slight
constrictions, measuring (11–)14–17(–21) x
5–7(–8) μm;
cosmopolitan...........................................................................................................
<italic><bold>M. tortile</bold></italic>9. Ascospores
longer..................................................................................................................................................................................
10</p></list-item><list-item><p>10. Ascospores 3-septate, elliptic-oblong, deeply constricted at the septa,
measuring 24–26 x 8–9 μm; North America.......... <italic><bold>M.
resinicola</bold></italic>10. Ascospores longer,
fusoid.......................................................................................................................................................................
11</p></list-item><list-item><p>11. Ascospores 3-septate, constricted at the median septum, measuring
27–33 x 7–8.5 μm; China and northwestern North
America.......................................................................................................................
<italic><bold>M. oblongisporum</bold></italic>11. Ascospores
longer..................................................................................................................................................................................
12</p></list-item><list-item><p>12. Ascospores 3-(4–5)-septate, measuring
(26–)30–34(–40) x (10–)12–13(–15) μm;
on <italic>Thuja</italic>, cosmopolitan........................ <italic><bold>M.
thujarum</bold></italic>12. Ascospores wider, 3–7(–9)-septate, with
swollen middle cells, 32–44 x 10–15 μm; on <italic>Austrocedrus
chilensis</italic>, Argentina
<italic>..............................................................................................................................................................................
<bold>M. andinense</bold></italic></p></list-item><list-item><p>13. Ascospores 5–7-septate, measuring 40–50 x 2–2.5
μm, slightly constricted at central septa; North America and
Europe..........................................................................................................................................
<italic><bold>M. scolecosporum</bold></italic>13. Ascospores longer, with more septa,
less
constricted............................................................................................................................
14</p></list-item><list-item><p>14. Ascospores 7–9(–11)-septate, measuring
(48–)54–62(–65) x 2.7–3 μm; North
America...................................................... <italic><bold>M.
parvulum</bold></italic>14. Ascospores (10–)11–14-septate, measuring
(54–)58–70(–75) x 3–4 μm; North
America....................................................... <italic><bold>M.
australe</bold></italic></p></list-item></list></p><p>Sequence data presented here (<xref ref-type="fig" rid="fig1">Fig.
1</xref>), based on an analysis of 10 of the 15 currently recognised
species (<xref ref-type="table" rid="tbl1">Table 1</xref>), do not
support subgenus <italic>Lophiopsis sensu</italic> Lohman
(<xref ref-type="bibr" rid="ref57">1932b</xref>): <italic>Mytilinidion
scolecosporum</italic> (<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=305.34&link_type=cbs">CBS
305.34</ext-link>) does not belong to the same clade as <italic>M. australe</italic>(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=301.34&link_type=cbs">CBS 301.34</ext-link>)
(<xref ref-type="fig" rid="fig1">Fig. 1</xref>). This implies that the
scolecospore has independently evolved at least twice within the family. Data
do however support the association of fusoid or spindle-shaped spores
belonging to <italic>M. thujarum</italic> (EB 0268 / BPI 879797) and to <italic>M.
andinense</italic> (<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123562&link_type=cbs">CBS
123562</ext-link> / BPI 878737), thus defining a lineage for this type of
spore within the genus. On the other hand, species possessing short, squat
phragmospores, namely <italic>M. acicola</italic> (EB 0349 / BPI 879794, EB 0379 / BPI
879793)<italic>, M. tortile</italic> (EB 0377 / BPI 879798), and <italic>M.
resinicola</italic> (<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=304.34&link_type=cbs">CBS
304.34</ext-link>) display complex relationships with species possessing
elongate phragmospores, such as <italic>M. californicum</italic> (EB 0385 / BPI
879795), <italic>M. mytilinellum</italic> (EB 0386 / BPI 879796,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=303.34&link_type=cbs">CBS 303.34</ext-link>) and
<italic>M. rhenanum</italic> (EB 0341,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=135.45&link_type=cbs">CBS 135.45</ext-link>). This
indicates that phragmospores with different length to width ratios have also
evolved multiple times within the genus
(<xref ref-type="fig" rid="fig1">Fig. 1</xref>). A manuscript
currently in preparation (Boehm <italic>et al.</italic>) will address speciation
events within the <italic>Mytilinidiaceae</italic>. Despite the lack of molecular
support for the subgenus <italic>Lophiopsis</italic>, it is included in the key below
to facilitate species identification.</p><p><italic><bold>Lophium</bold></italic> Fr., Syst. Mycol. 2: 534. 1823.</p><p><list list-type="simple"><list-item><p>= <italic>Lophidium</italic> P. Karst., Bidrag. Kännedom Finlands Natur Folk.
23: 33, 247. 1873.</p></list-item></list></p><p>The genus <italic>Lophium</italic> is characterised by fragile, conchate ascocarps,
sometimes seated on a foot-like base or sessile directly on the substrate. The
thin-walled scleroparenchymatous peridium encloses a basal hamathecium of
narrow trabeculate pseudoparaphyses, with very elongate asci, each bearing one
fascicle of transversely septate filiform ascospores, often spirally arranged.
The type species, <italic>Lophium mytilinum</italic>(<xref ref-type="fig" rid="fig11">Fig. 11J–K</xref>), is
cosmopolitan in the temperate zones and has been recorded from both sides of
the Atlantic (<xref ref-type="bibr" rid="ref122">Zogg 1962</xref>,
<xref ref-type="bibr" rid="ref8">Barr 1990a</xref>). Zogg
(<xref ref-type="bibr" rid="ref122">1962</xref>) described two
additional species, namely <italic>L. elegans</italic> on <italic>Juniperus</italic> from
alpine regions of France, Italy and Switzerland, and <italic>L. mayorii</italic> on
<italic>Pinus</italic> and <italic>Larix</italic> from the European Alps. Like
<italic>Mytilinidion</italic>, most species of <italic>Lophium</italic> have only been
recovered from coniferous substrates. The exception being the recently
described <italic>L. igoschinae</italic>, recovered on <italic>Dryas octopetala</italic> and
<italic>D. crenulata</italic> (<italic>Rosaceae</italic>) from Russia and Greenland
(<xref ref-type="bibr" rid="ref23">Chlebicki & Knudsen
2001</xref>).</p><p>Three isolates of the type species, <italic>L. mytilinum</italic>, were surveyed
(<xref ref-type="table" rid="tbl1">Table 1</xref>), two from the United
States, one from Michigan (<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=269.34&link_type=cbs">CBS
269.34</ext-link>) and one from New York
(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123344&link_type=cbs">CBS 123344</ext-link> / BPI
878736), and one from Sweden
(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=114111&link_type=cbs">CBS 114111</ext-link>). An
additional species of <italic>Lophium</italic>, namely a single-spored isolate of
<italic>L. elegans</italic> from France (EB 0366 / BPI 879792), was included as well
(<xref ref-type="table" rid="tbl1">Table 1</xref>). Both species are
morphologically similar, with <italic>L. elegans</italic> having spirally arranged
spores in the ascus and <italic>L. mytilinum</italic> having them in parallel
orientation (<xref ref-type="bibr" rid="ref122">Zogg 1962</xref>).
Molecular data indicate that the two species are not closely related within
the family. <italic>Lophium mytilinum</italic>, with filiform ascospores, shows a
close phylogenetic relationship however to the genus <italic>Quasiconcha</italic> (EB
QR), with reticulated didymospores (<xref ref-type="fig" rid="fig1">Fig.
1</xref>). Although having dissimilar spores, the fruitbodies of both
taxa are remarkably similar in their shape, size and fragility.</p></sec><sec><title>Key to the species of <italic>Lophium</italic></title><p><list list-type="simple"><list-item><p>1. Fruitbody erect, conchate, with thin-walled sclerenchymatoid
peridium...................................................................................................
2
1. Fruitbody conchate, but crowded, band- or ridge-like, horizontal to
recumbent and elongated; ascospores arranged parallel in the ascus, measuring
(60–)80–100(–110) x 3–4(–5) μm; Europe,
Russian Far East............. <italic><bold>L. mayorii</bold>Note</italic>: Transferred
to the genus <italic>Zoggium</italic> (<xref ref-type="bibr" rid="ref117">Vasilyeva
2001</xref>).</p></list-item><list-item><p>2. Ascospores filiform, 12–15-septate, measuring 78–86 x
2.6–3 μm; on <italic>Dryas</italic>, Greenland,
Russia.................................... <italic><bold>L. igoschinae</bold></italic>2.
Ascospores filiform, but longer; on
conifers..............................................................................................................................................
3</p></list-item><list-item><p>3. Ascospores arranged parallel in the ascus; measuring
(130–)170–250(–300) x 1–2(–2.5) μm;
cosmopolitan..................... <italic><bold>L. mytilinum</bold></italic>3.
Ascospores spirally arranged in the ascus; measuring
(200–)260–280(–300) x 2 μm;
Europe............................................... <italic><bold>L.
elegans</bold></italic></p></list-item></list></p><p><italic><bold>Zoggium</bold></italic> Lar.N. Vassiljeva, Mikol. Fitopatol. 35: 17.
2001.</p><p>Zogg described <italic>Lophium mayorii</italic> on <italic>Pinus</italic> and
<italic>Larix</italic> from the Swiss and French Alps, but noted that it differed from
other species of <italic>Lophium</italic> in having rigid, band-forming ascomata, with
a less fragile peridium as compared to <italic>Lophium</italic> and
<italic>Mytilinidion</italic>. Vasilyeva
(<xref ref-type="bibr" rid="ref117">2001</xref>) found the same fungus
in the Russian Far East and stated that it differed sufficiently from other
species of <italic>Lophium</italic> in having gross, erumpent crowded ascomata, band-
or ridge-like in appearance, as compared to the smaller, fragile, and entirely
superficial fruitbodies typical of species of <italic>Lophium</italic> and made the
transfer to <italic>Zoggium mayorii</italic>. Molecular data are presently
lacking.</p><p><italic><bold>Ostreola</bold></italic> Darker, Canad. J. Bot. 41: 1383. 1963.</p><p>Barr (<xref ref-type="bibr" rid="ref4">1975</xref>,
<xref ref-type="bibr" rid="ref8">1990a</xref>) recognised two genera
with muriform ascospores in the <italic>Mytilinidiaceae</italic>, namely
<italic>Ostreichnion</italic> and <italic>Ostreola</italic>. Darker
(<xref ref-type="bibr" rid="ref26">1963</xref>) originally established
the genus <italic>Ostreola</italic> for dictyospored forms that otherwise resembled
species of <italic>Mytilinidion</italic> – that is, mytilinidioid counterparts
to <italic>Hysterographium sensu</italic> Zogg
(<xref ref-type="bibr" rid="ref122">1962</xref>). Barr
(<xref ref-type="bibr" rid="ref8">1990a</xref>) differentiated
<italic>Ostreola</italic> from <italic>Ostreichnion</italic> by smaller ascospores in the
former, and recognised two species from North America, <italic>Ot. consociata</italic>from northeastern North America, and <italic>Ot. formosa</italic>, the latter common
on conifers in western North America and Europe, with spores similar to those
of <italic>Hysterobrevium mori</italic>. Tilak & Kale
(<xref ref-type="bibr" rid="ref112">1968</xref>) added another two
species from India, namely <italic>Ot. indica</italic> and <italic>Ot. ziziphi</italic>,
surprisingly both from non-coniferous substrates. Molecular data are presently
lacking for this genus.</p></sec><sec><title>Key to the species of <italic>Ostreola</italic></title><p><list list-type="simple"><list-item><p>1. Ascomata on coniferous hosts; North America,
Europe...........................................................................................................................
2
1. Ascomata on non-coniferous hosts;
India.................................................................................................................................................
3</p></list-item><list-item><p>2. Base of ascoma foot-like, immersed in substrate; ascospores
3–5(–7)-septate, with a longitudinal septum in the mid-cells,
14–18(–22) x 5–7 μm; on <italic>Picea</italic>, Northeastern
North
America....................................................................
<italic><bold>Ot. consociata</bold></italic>2. Base of ascoma tapered or applanate on
surface of substrate; ascospores (3–)5(–6)-septate, wider than in
<italic>O. consociata</italic>, 15–21 x 6.5–9.5 μm; alpine, on
<italic>Abies</italic>, Europe and Western North
America................................................. <italic><bold>Ot.
formosa</bold></italic></p></list-item><list-item><p>3. Ascospores transversely 3–7-septate, with 2–3 longitudinal
septa, slightly constricted in the middle; 24–30 x 8–9.6 μm; on
culms of <italic>Maduca</italic>,
India.........................................................................................................................................................
<italic><bold>Ot. indica</bold></italic>3. Ascospores as above but smaller, 19–23
x 6–7.5 μm; on culms of <italic>Ziziphus</italic>,
India...................................................................
<italic><bold>Ot. ziziphi</bold></italic></p></list-item></list></p><p><italic><bold>Gloniaceae</bold></italic> (Corda) E.W.A. Boehm, C.L. Schoch & J.W.
Spatafora 2009, <italic>Pleosporomycetidae fam. incertae sedis.</italic></p><p><list list-type="simple"><list-item><p>= <italic>Gloniaceae</italic> Corda, Icon. Fung. (Abellini) 5: 34. 1842.</p></list-item></list></p><p>Corda (<xref ref-type="bibr" rid="ref25">1842</xref>) originally
proposed the <italic>Gloniaceae</italic> as an intrafamilial taxonomic rank under the
family <italic>Hysteriaceae</italic>, to comprise <italic>Hysterographium</italic> and
<italic>Glonium</italic>. Boehm <italic>et al.</italic>(<xref ref-type="bibr" rid="ref18">2009</xref>) emended and restricted
the circumscription and elevated the taxon to family rank. The genus
<italic>Glonium</italic> was retained as circumscribed first by von Höhnel
(<xref ref-type="bibr" rid="ref41">1918</xref>) and then by Petrak
(<xref ref-type="bibr" rid="ref86">1923a</xref>). We feel justified in
reinstating the <italic>Gloniaceae</italic> and, more importantly, recognising it at
family rank for a single genus, because of the high support the group receives
in a recent four-gene analysis (<xref ref-type="bibr" rid="ref18">Boehm
<italic>et al.</italic> 2009</xref>), and corroborated here.</p><p><italic><bold>Glonium</bold></italic> Muhl., Cont. Lab. Plant Disease Sci. Fac. Agric.
Gifu Univ. 101. 1813.</p><p><list list-type="simple"><list-item><p>= <italic>Solenarium</italic> Spreng., Syst. Veg. 4(1): 376, 414. 1827.</p></list-item><list-item><p>= <italic>Psiloglonium</italic> Höhn., Ann. Mycol. 16(1): 149. 1918.</p></list-item></list></p><p>The genus <italic>Glonium</italic> is characterised by modified hysterothecia,
progressively dichotomously branched, laterally anastomosed along their length
to form radiating flabelliform or pseudostellate composites, usually seated
upon a conspicuous brown felt-like subiculum, sometimes borne in a stroma
(<xref ref-type="bibr" rid="ref122">Zogg 1962</xref>). Hysterothecia in
vertical section globose to obovoid, typically with a thick, three-layered
peridium, but fragile, unlike the robust peridium of the
<italic>Hysteriaceae</italic>. Luttrell
(<xref ref-type="bibr" rid="ref68">1953</xref>) described the
development of the ascocarp in the type species, <italic>G. stellatum</italic> as
composed of small pseudoparenchymatous cells, the outer layer heavily
encrusted with pigment and often longitudinally striate on the surface, the
middle layer lighter in pigmentation and the inner layer distinctly
thin-walled, pallid and compressed. The hamathecium consisted of persistent
narrow cellular pseudoparaphyses, often borne in a gel matrix, with tips
darkened or branched at maturity. Bitunicate asci are borne in a basal layer
and at maturity are typically clavate to cylindric, bearing eight ascospores,
overlapping biseriate; ascospores ranging from hyaline to pale yellow,
1-septate, conspicuously constricted at the septum, fusoid in outline, with at
least one end, often both, acuminate, and showing bipolar asymmetry.</p><p>Zogg (<xref ref-type="bibr" rid="ref122">1962</xref>) listed three
species that he grouped together in his key, that later formed the basis for
the <italic>Gloniaceae</italic> (<xref ref-type="bibr" rid="ref18">Boehm <italic>et
al.</italic> 2009</xref>). These are the type species, <italic>G. stellatum</italic>(<xref ref-type="fig" rid="fig12">Fig. 12A–E</xref>), <italic>G.
graphicum</italic>, and <italic>G. compactum</italic>, the latter associated with both
subicula and stroma. To these three species, we can add the recently described
saxicolous, terricolous and lignicolous <italic>G. circumserpens</italic>(<xref ref-type="fig" rid="fig12">Fig. 12F–H</xref>) from
Tasmania (<xref ref-type="bibr" rid="ref43">Kantvilas & Coppins
1997</xref>). Although von Höhnel
(<xref ref-type="bibr" rid="ref41">1918</xref>) and Petrak
(<xref ref-type="bibr" rid="ref86">1923a</xref>) stressed the
importance of subiculum as a synapomorphic character state, Zogg
(<xref ref-type="bibr" rid="ref122">1962</xref>) noted that <italic>G.
graphicum</italic> may or may not be associated with a subiculum. This, combined
with the observation that <italic>P. lineare</italic> may also on occasion be
associated with subiculum, led Zogg not to accept the genus
<italic>Psiloglonium</italic>. Data presented here and elsewhere
(<xref ref-type="bibr" rid="ref18">Boehm <italic>et al.</italic> 2009</xref>),
however, indicate that the synapomorphic character state is not subicula
<italic>per se</italic>, but the ascomata, which are modified hysterothecia that are
progressively dichotomously branched, to form radiating pseudostellate
composites. This is most pronounced in <italic>G. stellatum</italic> and <italic>G.
circumserpens</italic>, but may also be found to a lesser extent in <italic>G.
graphicum</italic> (<xref ref-type="bibr" rid="ref122">Zogg 1962</xref>).
One distinguishing feature that separates <italic>G. stellatum</italic> from <italic>G.
circumserpens</italic> is that, although both are associated with subicula, in the
former this extends as a wide margin in front of the developing hysterothecia
(<xref ref-type="fig" rid="fig12">Fig. 12A–C</xref>), whereas in
<italic>G. circumserpens</italic> (<xref ref-type="fig" rid="fig12">Fig.
12F–G</xref>) the subicula is only associated with the under
surface of the hysterothecia, closely appressed to the substrate, with only
minor deviations from the long axis of the fruitbody.</p><p><fig position="float" id="fig12"><label>Fig. 12.</label><caption><p>The <italic>Gloniaceae</italic>. A–E. <italic>Glonium stellatum</italic> [ANM 41
(ILLS), U.S.A.; not incl.], arrows in 12B, subiculum. F–H. <italic>Glonium
circumserpens</italic> [<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123343&link_type=cbs">CBS
123343</ext-link> (BPI 878739), Tasmania]. Scale bar (habitat) = 1 mm; Scale
bar (spores and asci) = 10 μm.</p></caption><graphic xlink:href="49fig12"></graphic></fig></p></sec><sec><title>Key to the species of <italic>Glonium</italic></title><p><list list-type="simple"><list-item><p>1. Hysterothecia associated and seated upon a thin crust-like stroma, or
arising from within a stromal crust; stroma itself seated on subiculum;
didymospores spindle-shaped with the upper cell slightly swollen and larger
than the lower cell, measuring 24–28 x 5–6 μm;
Africa.....................................................................................................
<italic><bold>G. compactum</bold></italic>1. Hysterothecia not associated with
stroma................................................................................................................................................
2</p></list-item><list-item><p>2. Hysterothecia somewhat branched, irregular, “graphoid”;
without well-developed subiculum; didymospores oblong to spindle-shaped; upper
cell pear-shaped, constricted at septum; both ends acuminate, measuring
(13–)15–18(–21) x (3–)5–6 μm; on <italic>Pinus,
Juniperus</italic>,
Europe...........................................................................
<italic><bold>G. graphicum</bold></italic>2. Hysterothecia in mature specimens highly
bifurcated, closely appressed to the substrate, dichotomously branched to form
irregular creeping masses; usually seated upon or sitting behind a front of
well-developed brown to black
subiculum.................................................................................................................................
3</p></list-item><list-item><p>3. Didymospores hyaline, constricted at the septum, apices pointed,
measuring (15–)16–17 x 6–7 μm; on soil (terricolous) or
rock (saxicolous), or lignicolous;
Tasmania..............................................................................
<italic><bold>G. circumserpens</bold></italic>3. Didymospores oblong to spindle-shaped;
upper cell pear-shaped, constricted at septum; both ends acuminate, measuring
(18–)21–26(–28) x (4–)5–6(–7) μm;
cosmopolitan...............................................................................................
<italic><bold>G. stellatum</bold></italic></p></list-item></list></p><p>Four isolates were surveyed for the <italic>Gloniaceae</italic>. Two of <italic>G.
stellatum</italic>, from Michigan
(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=207.34&link_type=cbs">CBS 207.34</ext-link>) and
Tennessee (ANM 32), the United States, and two of <italic>G. circumserpens</italic>,
recently isolated from wood
(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123342&link_type=cbs">CBS 123342</ext-link> / BPI
878738) and dolerite stone (<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123343&link_type=cbs">CBS
123343</ext-link> / BPI 878739) from Tasmania
(<xref ref-type="table" rid="tbl1">Table 1</xref>). Molecular data
indicate that all four isolates are closely related. Surprisingly, this clade
also includes multiple isolates of <italic>Cenococcum geophilum</italic>, an
ecologically important ectomycorrhizal fungus with a global distribution and a
wide host range, but with no known teleomorph
(<xref ref-type="bibr" rid="ref54">LoBuglio <italic>et al.</italic>1996</xref>).</p><p><italic><bold>Farlowiella</bold></italic> Sacc., Syll. Fung. 9: 1101. 1891.</p><p><list list-type="simple"><list-item><p>= <italic>Farlowia</italic> Sacc., Syll. Fung. 2: 727. 1883.</p></list-item></list></p><p>Recent molecular data (<xref ref-type="bibr" rid="ref100">Schoch <italic>et
al.</italic> 2006</xref>; <xref ref-type="bibr" rid="ref18">Boehm <italic>et
al.</italic> 2009</xref>) support the transference of the genus
<italic>Farlowiella</italic> from the <italic>Hysteriaceae</italic>, and its current placement
as <italic>Pleosporomycetidae gen. incertae sedis</italic>. The genus is characterised
by 1-celled pedicellate slightly laterally compressed amerospores, the upper
cell pigmented and much larger than the lower, which remains hyaline or
moderately pigmented, and can be considered as an associated papilla. The
hysterothecia are somewhat laterally compressed, but nonetheless thick-walled
and with a prominent sunken slit. They can be solitary to gregarious, but
remain erect, and elevated, presenting an almost stipitate appearance.
Anamorphs have been described in the genus <italic>Acrogenospora</italic>(<xref ref-type="bibr" rid="ref38">Goh <italic>et al.</italic> 1998</xref>).
Two species are recognised, namely <italic>Farlowiella carmichaeliana</italic> from
Europe (Belgium, England, Germany, Switzerland), from the bark and wood of
<italic>Fagus, Quercus, Sorbus</italic> and <italic>Prunus</italic>, and <italic>F. australis</italic>known only from the original collection on <italic>Phylica arborea</italic> from
Tristan da Cunha in the South Atlantic
(<xref ref-type="bibr" rid="ref29">Dennis 1955</xref>). Sequence data
from two isolates of <italic>F. carmichaeliana</italic>(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=206.36&link_type=cbs">CBS 206.36</ext-link> and
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=179.73&link_type=cbs">CBS 179.73</ext-link>)
indicate that this taxon lies quite distant from the <italic>Hysteriaceae</italic>(<xref ref-type="fig" rid="fig1">Fig. 1</xref>). An additional isolate
of the anamorph, <italic>Acrogenospora sphaerocephala</italic>(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=164.76&link_type=cbs">CBS 164.76</ext-link>),
further supports the current placement of the genus <italic>Farlowiella</italic>.</p></sec><sec><title>Key to the species of <italic>Farlowiella</italic></title><p><list list-type="simple"><list-item><p>1. Ascospores unequally 2-celled; upper cell pigmented, much larger than
the lower cell, which is smaller and hyaline, together measuring 18–21 x
7–12 μm;
Europe.............................................................................................................
<italic><bold>F. carmichaeliana</bold></italic>1. Ascospores as above, but smaller,
13–15 x 6–7.5 μm; Tristan da
Cunha...............................................................................
<italic><bold>F. australis</bold></italic></p></list-item></list></p></sec></sec><sec><title>CONCLUSIONS</title><p>Hysteriaceous fungi are an ancient and ecologically successful group of
organisms, as attested by their wide geographic distribution on a multitude of
gymnosperm and angiosperm host species. Whereas the <italic>Mytilinidiaceae</italic>are found almost exclusively on conifers, the <italic>Hysteriaceae</italic> occur
primarily on angiosperms (<xref ref-type="bibr" rid="ref122">Zogg
1962</xref>). Presumably, the <italic>Hysteriaceae</italic> underwent rapid
speciation in response to the angiosperm radiation of the mid- to
late-Cretaceous, 65–100 mya (<xref ref-type="bibr" rid="ref84">Palmer
<italic>et al.</italic> 2004</xref>). However, this must have occurred prior to
the complete loss of continental contiguity, which occurred during the same
time period. This is because we see today a remarkable degree of intraspecific
stability, in both morphology and sequence data, among geographically
disparate accessions (<xref ref-type="fig" rid="fig1">Fig. 1</xref>).
For example, little morphological or sequence variation was detected in
<italic>Hysterium angustatum</italic>, from North America
(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123334&link_type=cbs">CBS 123334</ext-link>), Kenya
(GKM 243A), New Zealand (SMH 5211.0), and South Africa (CMW 20409;
<xref ref-type="bibr" rid="ref49">Lee & Crous 2003</xref>).
Similarly, little variation was detected in <italic>Psiloglonium clavisporum</italic>,
from Kenya (GKM L172A, GKM 344A) and North America (<italic>e.g.</italic>,
<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123338&link_type=cbs">CBS 123338</ext-link>), or in
<italic>Oedohysterium sinense</italic>, from South Africa
(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123345&link_type=cbs">CBS 123345</ext-link>) and
North America (EB 0339). As we are presumably sampling remnants of once
contiguous sexual populations, their similarity today must imply that
speciation occurred prior to complete genetic isolation. The break-up of
Pangea during the Triassic 200 mya, and the formation of the nascent central
Atlantic Ocean, separating Gondwana from Laurasia, during the Jurassic, 150
mya, must have effectively disrupted once contiguous populations. Although
most flowering plant families were established by the end of the Cretaceous,
65–70 mya, it is now believed that they diversified into their present
lineages (<italic>e.g.</italic>, eudicots, Magnoliids and monocots) much earlier,
around 140 mya (<xref ref-type="bibr" rid="ref27">Davies <italic>et al.</italic>2004</xref>, <xref ref-type="bibr" rid="ref84">Palmer <italic>et al.</italic>2004</xref>, <xref ref-type="bibr" rid="ref80">Moore <italic>et al.</italic>2007</xref>). This may have allowed for remnants of once contiguous
populations to colonise early angiosperm lineages, prior to the complete
dissolution of continental integrity during the mid- to late-Cretaceous.
Recent studies (<xref ref-type="bibr" rid="ref64">Lücking <italic>et
al.</italic> 2009</xref>), based on a recalibration of published molecular
clock trees, using internally unconstrained, uniform calibration points, have
suggested an origin for the fungi between 760 mya to 1.0 bya, with the origin
of the <italic>Ascomycota</italic> set at 500-650 mya. Whatever the timing,
hysteriaceous fungi incurred little appreciable intraspecific morphological or
genetic (<italic>e.g.</italic>, nuLSU, nuSSU, <italic>TEF1</italic> and <italic>RPB2</italic>) change
over significant periods of geologic time, on different continents. Thus, with
the exception of <italic>Hb. mori</italic>, and perhaps, <italic>Gp. subrugosa</italic> (see
below), most members of the <italic>Hysteriaceae</italic> appear to be stable
species.</p><p>Sequence data indicate that <italic>Hb. mori</italic> occurs in both Clades A and
D. However, analysis of <italic>Hb. mori</italic> specimens originating from each
clade (<italic>e.g.</italic>, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123563&link_type=cbs">CBS
123563</ext-link> / BPI 878731, and others, in Clade A <italic>versus</italic> GKM
1013 / BPI 879788 in Clade D), failed to find any appreciable difference in
either spore morphology (<italic>e.g.</italic>, septation, pigmentation, symmetry, or
measurement), substrate-choice, or features associated with the
hysterothecium. Likewise, no morphological difference could be detected among
genetically unrelated accessions of <italic>Gp. subrugosa</italic>, from South Africa
(<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123346&link_type=cbs">CBS 123346</ext-link> / BPI
878735), in Clade D, <italic>versus</italic> those from Kenya (GKM 1214 / BPI 879776)
and Cuba (SMH 557 / BPI 879777), outside of Clade D. These two examples
illustrate a lack of correspondence between the morphospecies concept
(<xref ref-type="bibr" rid="ref19">Burnett 2003</xref>) and the
genealogical concordance phylogenetic species recognition concept
(<xref ref-type="bibr" rid="ref110">Taylor <italic>et al.</italic>2000</xref>), the latter indicating here the presence of cryptic species
within the two morphospecies. <italic>Hysterobrevium mori</italic> and, to a lesser
extent, <italic>Gp. subrugosa</italic>, may represent examples of convergent
evolution, whereby similar ascospores borne in hysterothecia have evolved
multiple times within the family. This is supported by the polyphyly inherent
in the circumspection of the classical genera of the <italic>Hysteriaceae</italic>(<italic>e.g., Gloniopsis, Glonium, Hysterium</italic>, and <italic>Hysterographium</italic>),
revealed by recent studies (<xref ref-type="bibr" rid="ref100">Schoch <italic>et
al.</italic> 2006</xref>, <xref ref-type="bibr" rid="ref18">Boehm <italic>et
al.</italic> 2009</xref>, <xref ref-type="bibr" rid="ref81">Mugambi &
Huhndorf 2009</xref>). Alternatively, <italic>Hb. mori</italic> and <italic>Gp.
subrugosa</italic> may have retained ancestral character states, and thus may
represent evolutionary lineages that did not incur appreciable morphological
change, while at the same time accumulating sufficient genetic change to fall,
in the case of <italic>Hb. mori</italic>, into distant clades within the family. If
this is the case, then these two taxa may represent examples of speciation in
progress, with genetic change preceding morphological change, thus differing
from independent convergent character states. Whatever the mechanism, it is
difficult to see how <italic>Hb. mori</italic>, for example, may be classified into
different species, in different genera (<italic>e.g., Hysterobrevium</italic> and
<italic>Oedohysterium</italic>), without a sound morphological basis. We conclude that
both <italic>Hb. mori</italic> and <italic>Gp. subrugosa</italic> contain genetically
unrelated, cryptic, and potentially different biological species, that can not
at present be morphologically differentiated.</p><p>Although there are examples of concordance between morphological and
molecular data in this study (see below), these are few. For the most part,
molecular data support the premise of a large number of convergent
evolutionary lineages, sharing similar spore morphologies, but that are not
closely related. This resulted in a polyphyletic core set of genera for the
<italic>Hysteriaceae</italic>, and presented us with a complicated picture of past
speciation events within the family (<xref ref-type="bibr" rid="ref18">Boehm
<italic>et al.</italic> 2009</xref>). To achieve a natural phylogeny, that is,
one based on the concordance of morphological and molecular data, required
that we break-up what were once thought to be stable genera. Thus, two species
of <italic>Hysterium</italic> were transferred to <italic>Oedohysterium</italic> (<italic>Od.
insidens</italic> and <italic>Od. sinense</italic>), and two species of
<italic>Gloniopsis</italic> to <italic>Hysterobrevium</italic> (<italic>Hb. smilacis</italic> and
<italic>Hb. constrictum</italic>). While <italic>Hysterographium</italic>, with the type
<italic>Hg. fraxini</italic>, was removed from the <italic>Hysteriaceae</italic>(<xref ref-type="bibr" rid="ref18">Boehm <italic>et al.</italic> 2009</xref>),
some of its species remained within the family, transferred here to
<italic>Oedohysterium</italic> (<italic>Od. pulchrum</italic>), <italic>Hysterobrevium</italic>(<italic>Hb. mori</italic>) and <italic>Gloniopsis</italic> (<italic>Gp. subrugosa</italic>). New
species were described (<italic>e.g., Gp. arciformis</italic> and <italic>Gp.
kenyensis</italic>) which would previously have been classified in
<italic>Hysterographium</italic>, but are now accommodated in <italic>Gloniopsis</italic>.
Molecular data necessitated that both <italic>Gloniopsis</italic> and
<italic>Hysterobrevium</italic> include hyaline and pigmented dictyospores, and the
genus <italic>Oedohysterium</italic>, both phragmospores and dictyospores. This, then,
de-emphasised spore morphology as a synapomorphic character state. Likewise,
the genus <italic>Glonium sensu</italic> Zogg
(<xref ref-type="bibr" rid="ref122">1962</xref>) was divided into
<italic>Psiloglonium</italic> in the <italic>Hysteriaceae</italic> and <italic>Glonium</italic> in the
<italic>Gloniaceae</italic> (<xref ref-type="bibr" rid="ref18">Boehm <italic>et al.</italic>2009</xref>), and, more recently, <italic>Anteaglonium</italic> in the
<italic>Pleosporales</italic> (<xref ref-type="bibr" rid="ref81">Mugambi &
Huhndorf 2009</xref>). These taxonomic changes were unexpected, as they
were not premised on past assumptions of synapomorphy related to spore
morphology (<xref ref-type="bibr" rid="ref122">Zogg 1962</xref>).
Although we have included here a total of 59 accessions, representing 22
species in seven genera, for the <italic>Hysteriaceae</italic>, and another 62 outside
of the family (<xref ref-type="table" rid="tbl1">Table 1</xref>), taxon
sampling may still be insufficient. Clearly, additional species and genera
need to be sampled before a complete picture emerges for the family.</p><p>The quest for synapomorphic character states that correlate with molecular
data was one of the goals of this study. If traditional character states
associated with spore septation/pigmentation or the fruitbody
(<xref ref-type="bibr" rid="ref122">Zogg 1962</xref>) can not be relied
upon to deduce phylogeny, are there other character states that can be
emphasised instead? Two examples are discussed below, the first relating to
spore morphology, the second to characters associated with the fruitbody.
Although both <italic>Oedohysterium</italic> and <italic>Hysterium</italic> possess similar
pigmented asymmetric phragmospores, species of <italic>Oedohysterium</italic> can be
morphologically differentiated by the possession of an enlarged supra-median
cell. Molecular data also revealed that a species previously classified as
<italic>Hysterographium</italic>, namely <italic>Hg. pulchrum</italic>, belonged to
<italic>Oedohysterium</italic>, despite the presence of dictyospores. Closer
inspection, however, reveals that the dictyospores of <italic>Od. pulchrum</italic>also possess a swollen supra-median cell. Additionally, a certain number of
spores remain as transversely septate phragmospores
(<xref ref-type="bibr" rid="ref21">Checa <italic>et al.</italic> 2007</xref>),
thus reinforcing its placement within <italic>Oedohysterium</italic>, and perhaps
underscoring the plasticity of spore septation configurations for this group
of fungi.</p><p>The second example relates to character states associated with the
fruitbody. Fruitbody morphology clearly supports the transfer of the genus
<italic>Glonium</italic> out of the <italic>Hysteriaceae</italic> to its own family, the
<italic>Gloniaceae</italic>, closely allied with the <italic>Mytilinidiales</italic>. The
<italic>Gloniaceae</italic> possess a modified hysterothecium, one in which the
frutibodies frequently bifurcate to a greater (<italic>e.g., G. stellatum</italic> and
<italic>G. circumserpens</italic>) or lesser (<italic>e.g., G. graphicum</italic>) degree, the
former two species with radiating stellate composites, usually seated on
subicula. This is in contrast to hysterothecia found in the
<italic>Hysteriaceae</italic> which may be gregarious, but are never laterally
anastomosed to form radiating composites. Additionally, the morphology of the
dehiscence slit found in the <italic>Gloniaceae</italic> is unlike that found in the
<italic>Hysteriaceae</italic>. In the <italic>Gloniaceae</italic>, the aperture is in most
cases evaginated, forming a miniscule crest, similar to the more extended
version found in some species in the <italic>Mytilinidiaceae</italic>; whereas, in the
<italic>Hysteriaceae</italic>, hysterothecia have deeply invaginated slits. Also,
hysterothecia found in the <italic>Gloniaceae</italic>, like those in the
<italic>Mytilinidiaceae</italic>, are considerably more fragile, as compared to those
found within the <italic>Hysteriaceae</italic>. These character states were either not
noted before (<italic>e.g.</italic>, swollen supra-median cell in
<italic>Oedohysterium</italic> and evaginated slit in <italic>Glonium</italic>), or were
noticed, but given less taxonomic weight (<italic>e.g.</italic>, modified
hysterothecium in <italic>Glonium</italic>; <xref ref-type="bibr" rid="ref122">Zogg
1962</xref>). These examples illustrate that morphological features can
be found that correlate with molecular data, despite the anomalies presented
by <italic>Hb. mori</italic> and <italic>Gp. subrugosa</italic>, mentioned earlier.</p><p>The hysterothecium, thick-walled, navicular, and with a prominent
longitudinal slit, has long been considered synapomorphic, defining the
<italic>Hysteriales</italic>. However, this type of fruitbody has evolved convergently
no less than five times within the <italic>Pleosporomycetidae</italic> (<italic>e.g.,
Farlowiella, Glonium, Anteaglonium, Hysterographium</italic> and the
<italic>Hysteriaceae</italic>). Similarly, thin-walled mytilinidioid (<italic>e.g.,
Ostreichnion</italic>) and patellarioid (<italic>e.g., Rhytidhysteron</italic>) ascomata
have also evolved at least twice within the subclass, the genera having been
transferred from the <italic>Mytilinidiaceae</italic> and <italic>Patellariaceae</italic>,
respectively, to the <italic>Hysteriaceae</italic>. As such, character states relating
not only to the external features of the ascoma, but to the centrum as well
(<italic>e.g.</italic>, cellular pseudoparaphyses <italic>versus</italic> trabeculae, etc.),
previously considered to represent synapomorphies among these fungi, in fact,
represent symplesiomorphies, and most likely have arisen multiple times
through convergent evolutionary processes in response to common selective
pressures. Similar findings have emerged for a number of other ascomycete
lineages within the <italic>Pezizomycotina</italic> (<italic>e.g.</italic>,
<xref ref-type="bibr" rid="ref101">Schoch <italic>et al.</italic>2009b</xref>). One selective advantage of the hysterothecium may be spore
discharge over prolonged periods of time, since some, if not most, species may
be perennial (Lohman <xref ref-type="bibr" rid="ref55">1931</xref>,
<xref ref-type="bibr" rid="ref58">1933a</xref>). The thick-walled
peridium further contributes to xerotolerance, as many of these fungi persist
on decorticated, weathered woody substrates prone to prolonged periods of
desiccation. Thus, the ability to perennate, and time spore discharge with
environmental conditions suitable for germination, spanning multiple seasons,
may be the driving force behind the repeated evolution of the
hysterothecium.</p></sec></body><back><ack><p>The authors wish to thank Alain Gardiennet (Veronnes, France), Gintaras
Kantvilas (Tasmanian Herbarium, Hobart, Tasmania), Marieka Gryzenhout (Dept.
Microbiology and Plant Pathology, Forestry and Agricultural Biotechnology
Institute, University of Pretoria, South Africa), Maria Inéz Messuti
and Laura Emma Lorenzo (Departamento de Botanica, Centro Regional
Universitario Bariloche, Universidad Nacional del Comahue, Quintral,
Bariloche, Rio Negro, Argentina), Eunice Nkansah (Kean University, Union, NJ,
U.S.A.), and Meredith Blackwell (Dept. Biological Sciences, Louisiana State
University, Baton Rouge, Louisiana, U.S.A.) for kindly supplying some of the
isolates used in this study (<xref ref-type="table" rid="tbl1">Table
1</xref>). The authors wish to thank Walter Gams (Baarn, The Netherlands)
for the Latin translations, and for his numerous helpful insights into the
taxonomic and nomenclatural issues raised by this work. We also wish to
acknowledge Scott Redhead (National Mycological Herbarium, Agriculture and
Agri-Food Canada, Ottawa, Canada) who provided helpful suggestions on the
manuscript prior to submission. E.W.A. Boehm wishes to acknowledge support
from the National Science Foundation (NSF) Major Research Instrumentation
Grant DBI 0922603. A.N. Miller acknowledges funding from the NSF through a
BS&I award (DEB0515558) and from Discover Life in America (DLIA 2005-15).
Part of this work was also funded by a grant from NSF (DEB-0717476) to J.W.
Spatafora (and C.L. Schoch until 2008). Work performed by C.L. Schoch after
2008 was supported in part by the Intramural Research Program of the NIH,
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