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<document id="00EA05F397E94E90934E1BECF053A5DE" ID-CLB-Dataset="6312" ID-DOI="10.11646/phytotaxa.9.1.13" ID-GBIF-Dataset="21a2a91a-28ae-40c8-911e-13d0c279e095" ID-ISSN="1179-3163" ID-Zenodo-Dep="4895360" IM.metadata_approvedBy="felipe" IM.taxonomicNames_approvedBy="felipe" checkinTime="1621604093317" checkinUser="felipe" docAuthor="Pressel, Silvia, Bidartondo, Martin I., Ligrone, Roberto &amp; Duckett, Jeffrey G." docDate="2010" docId="03F587D3FFDCB94DB5974E12FEFEFD69" docLanguage="en" docName="Phytotaxa.9.238-253.pdf" docOrigin="Phytotaxa 9" docStyle="DocumentStyle:E765CC7F3D29AC31B7D5F874DD99C2DD.4:Phytotaxa.2009-2010.journal_article" docStyleId="E765CC7F3D29AC31B7D5F874DD99C2DD" docStyleName="Phytotaxa.2009-2010.journal_article" docStyleVersion="4" docTitle="Haplomitrium" docType="treatment" docVersion="8" lastPageNumber="241" masterDocId="FFCCFFABFFDDB94EB500481FFFC0FFEB" masterDocTitle="Fungal symbioses in bryophytes: New insights in the Twenty First Century" masterLastPageNumber="253" masterPageNumber="238" pageNumber="239" updateTime="1698941776982" updateUser="ExternalLinkService" zenodo-license-document="CLOSED">
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<mods:title id="E1A4D3C7D52780E619B159B1A2439F55">Fungal symbioses in bryophytes: New insights in the Twenty First Century</mods:title>
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<mods:namePart id="81F47D7DA2866455D808542951EE2227">Pressel, Silvia</mods:namePart>
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<mods:namePart id="09ACC9442FB5D6E5B3F896EAC13EF673">Bidartondo, Martin I.</mods:namePart>
<mods:affiliation id="81148EBFFB38A9319074212E1A3E19F1">Imperial College London and Royal Botanic Gardens, Kew TW 9 3 DS, UK; email: m. bidartondo @ imperial. ac. uk</mods:affiliation>
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<mods:namePart id="926A1A0005D37E72D7065DB4DE1E6C33">Ligrone, Roberto</mods:namePart>
<mods:affiliation id="6421632DA33CFC4675DF3096A779A6FA">Dipartimento di Scienze ambientali, Seconda Università di Napoli, via Vivaldi 43, 81100 Caserta, Italy;</mods:affiliation>
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<mods:namePart id="19D851CF733C54C8213D534AD64E51AD">Duckett, Jeffrey G.</mods:namePart>
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<mods:title id="5CF80363B38119D1109F22009D04B99F">Phytotaxa</mods:title>
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<mods:date id="6F78984F2C5A382F7198A705EDAF2465">2010</mods:date>
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<treatment id="03F587D3FFDCB94DB5974E12FEFEFD69" ID-DOI="http://doi.org/10.5281/zenodo.4895362" ID-GBIF-Taxon="182486520" ID-Zenodo-Dep="4895362" LSID="urn:lsid:plazi:treatment:03F587D3FFDCB94DB5974E12FEFEFD69" httpUri="http://treatment.plazi.org/id/03F587D3FFDCB94DB5974E12FEFEFD69" lastPageId="3" lastPageNumber="241" pageId="1" pageNumber="239">
<subSubSection id="C346654EFFDCB94FB5974E12FA43F903" pageId="1" pageNumber="239" type="nomenclature">
<paragraph id="8BE336C5FFDCB94FB5974E12FE3FF9C2" blockId="1.[151,511,1549,1578]" box="[151,511,1549,1578]" pageId="1" pageNumber="239">
<heading id="D0AB81A9FFDCB94FB5974E12FE3FF9C2" bold="true" box="[151,511,1549,1578]" fontSize="12" level="1" pageId="1" pageNumber="239" reason="1">
<emphasis id="B928EAD7FFDCB94FB5974E12FE3FF9C2" bold="true" box="[151,511,1549,1578]" pageId="1" pageNumber="239">
<taxonomicName id="4C5C4D46FFDCB94FB5974E12FE93F9C2" authority="Dedecek" box="[151,339,1549,1577]" class="Haplomitriopsida" family="Haplomitriaceae" genus="Haplomitrium" kingdom="Plantae" order="Calobryales" pageId="1" pageNumber="239" phylum="Bryophyta" rank="genus">
<emphasis id="B928EAD7FFDCB94FB5974E12FE93F9C2" bold="true" box="[151,339,1549,1577]" italics="true" pageId="1" pageNumber="239">Haplomitrium</emphasis>
</taxonomicName>
and
<taxonomicName id="4C5C4D46FFDCB94FB4974E12FE3FF9C2" authority="K.I.Goebel" box="[407,511,1549,1577]" class="Haplomitriopsida" family="Treubiaceae" genus="Treubia" kingdom="Plantae" order="Treubiales" pageId="1" pageNumber="239" phylum="Bryophyta" rank="genus">
<emphasis id="B928EAD7FFDCB94FB4974E12FE3FF9C2" bold="true" box="[407,511,1549,1577]" italics="true" pageId="1" pageNumber="239">Treubia</emphasis>
</taxonomicName>
</emphasis>
</heading>
</paragraph>
<paragraph id="8BE336C5FFDCB94FB5974E45FA43F903" blockId="1.[151,1436,1626,2000]" pageId="1" pageNumber="239">
Until the recent and detailed electron microscope studies by
<bibRefCitation id="EFCD4B34FFDCB94FB66E4E45FB99F99F" author="Carafa, A. &amp; Duckett, J. G. &amp; Ligrone, R." box="[878,1113,1626,1652]" pageId="1" pageNumber="239" pagination="185 - 197" refId="ref7167" refString="Carafa, A., Duckett, J. G. &amp; Ligrone, R. (2003) Subterranean gametophytic axes in the primitive liverwort Haplomitrium harbour a unique type of endophytic association with aseptate fungi. New Phytologist 160: 185 - 197." type="journal article" year="2003">
Carafa
<emphasis id="B928EAD7FFDCB94FB6C54E45FBC2F998" box="[965,1026,1626,1651]" italics="true" pageId="1" pageNumber="239">et al.</emphasis>
(2003)
</bibRefCitation>
and
<bibRefCitation id="EFCD4B34FFDCB94FB1954E45FA5CF99F" author="Duckett, J. G. &amp; Carafa, A. &amp; Ligrone, R." box="[1173,1436,1626,1652]" pageId="1" pageNumber="239" pagination="797 - 813" refId="ref7597" refString="Duckett, J. G., Carafa, A. &amp; Ligrone, R. (2006 a) A highly differentiated glomeromycotean association with the mucilagesecreting, primitive antipodean liverwort Treubia: clues to the origins of mycorrhizas. American Journal of Botany 93: 797 - 813." type="journal article" year="2006">
Duckett
<emphasis id="B928EAD7FFDCB94FB1FB4E45FAF7F998" box="[1275,1335,1626,1651]" italics="true" pageId="1" pageNumber="239">et al.</emphasis>
(2006a)
</bibRefCitation>
our knowledge of the endophytes in these two genera was limited to early light microscope investigations (
<bibRefCitation id="EFCD4B34FFDCB94FB59F4EB8FE84F92A" author="Goebel, K." box="[159,324,1703,1729]" pageId="1" pageNumber="239" pagination="1 - 11" refId="ref7931" refString="Goebel, K. (1891) Morphologische und Biologische Studien. IV. Uber javanische Lebermoose. 1. Treubia. Annales de Jardin Botanique de Buitenzorg 9: 1 - 11." type="journal article" year="1891">Goebel 1891</bibRefCitation>
;
<bibRefCitation id="EFCD4B34FFDCB94FB4534EB8FE1CF92A" author="Grun, C." box="[339,476,1703,1729]" pageId="1" pageNumber="239" pagination="331 - 392" refId="ref8149" refString="Grun, C. (1914) Monographische Studien an Treubia insignis Goebel. Flora 106: 331 - 392." type="journal article" year="1914">Grun 1914</bibRefCitation>
; Lilienfeld 1911). These suggested that fungal associations in
<taxonomicName id="4C5C4D46FFDCB94FB1ED4EB8FF3DF903" authority="Dedecek" authorityName="Dedecek" class="Haplomitriopsida" family="Haplomitriaceae" genus="Haplomitrium" kingdom="Plantae" order="Calobryales" pageId="1" pageNumber="239" phylum="Bryophyta" rank="genus">
<emphasis id="B928EAD7FFDCB94FB1ED4EB8FA5BF92B" box="[1261,1435,1703,1728]" italics="true" pageId="1" pageNumber="239">Haplomitrium</emphasis>
Dedecek
</taxonomicName>
and
<taxonomicName id="4C5C4D46FFDCB94FB4364ED1FE50F90C" authority="K.I.Goebel" box="[310,400,1742,1767]" class="Haplomitriopsida" family="Treubiaceae" genus="Treubia" kingdom="Plantae" order="Treubiales" pageId="1" pageNumber="239" phylum="Bryophyta" rank="genus">
<emphasis id="B928EAD7FFDCB94FB4364ED1FE50F90C" box="[310,400,1742,1767]" italics="true" pageId="1" pageNumber="239">Treubia</emphasis>
</taxonomicName>
might be different from those found elsewhere in land plants and so this proved to be.
</paragraph>
</subSubSection>
<subSubSection id="C346654EFFDCB94DB5C64EEBFEFEFD69" lastPageId="3" lastPageNumber="241" pageId="1" pageNumber="239" type="description">
<paragraph id="8BE336C5FFDCB94CB5C64EEBFB3BFECD" blockId="1.[151,1436,1626,2000]" lastBlockId="2.[151,1436,152,294]" lastPageId="2" lastPageNumber="240" pageId="1" pageNumber="239">
Electron microscopy confirmed the presence of both extra and intracellullar hyphae in
<taxonomicName id="4C5C4D46FFDCB94FB1D74EEBFAF1F8E6" box="[1239,1329,1780,1805]" class="Haplomitriopsida" family="Treubiaceae" genus="Treubia" kingdom="Plantae" order="Treubiales" pageId="1" pageNumber="239" phylum="Bryophyta" rank="genus">
<emphasis id="B928EAD7FFDCB94FB1D74EEBFAF1F8E6" box="[1239,1329,1780,1805]" italics="true" pageId="1" pageNumber="239">Treubia</emphasis>
</taxonomicName>
(
<figureCitation id="13672A40FFDCB94FB0434EEBFA4AF8E5" box="[1347,1418,1780,1806]" captionStart="FIGURE 1" captionStartId="2.[151,259,1813,1837]" captionTargetBox="[155,1432,349,1790]" captionTargetId="figure-174@2.[155,1433,349,1790]" captionTargetPageId="2" captionText="FIGURE 1. The highly differentiated endophyte in Treubia pygmaea R.M.Schust.. A, B, D. Scanning electron micrographs. C, E, F. Transmission electron micrographs. A. Profile of thallus showing the intercellular fungal zone (ef) and the intracellular zone below it (if). B, C. Details of the intracellular fungus showing hyphal coils (h), fungal lumps (l) and a degenerating lump (dl). D, E. Intracellular fungus showing pseudoparenchymatous hyphae (arrowed). F. Fungal spores (s) with multilayered walls in the intercellular mucilage (*). Scale bars = (A) 200 µm; (B, D) 20 µm; (C, E, F) 5 µm." figureDoi="http://doi.org/10.5281/zenodo.4895364" httpUri="https://zenodo.org/record/4895364/files/figure.png" pageId="1" pageNumber="239">Fig. 1</figureCitation>
). The former forms pseudoparenchyma-like structures within the mucilage-filled intercellular spaces in the thalli (
<figureCitation id="13672A40FFDCB94FB5E34F5DFEA1F8B7" box="[227,353,1858,1884]" captionStart="FIGURE 1" captionStartId="2.[151,259,1813,1837]" captionTargetBox="[155,1432,349,1790]" captionTargetId="figure-174@2.[155,1433,349,1790]" captionTargetPageId="2" captionText="FIGURE 1. The highly differentiated endophyte in Treubia pygmaea R.M.Schust.. A, B, D. Scanning electron micrographs. C, E, F. Transmission electron micrographs. A. Profile of thallus showing the intercellular fungal zone (ef) and the intracellular zone below it (if). B, C. Details of the intracellular fungus showing hyphal coils (h), fungal lumps (l) and a degenerating lump (dl). D, E. Intracellular fungus showing pseudoparenchymatous hyphae (arrowed). F. Fungal spores (s) with multilayered walls in the intercellular mucilage (*). Scale bars = (A) 200 µm; (B, D) 20 µm; (C, E, F) 5 µm." figureDoi="http://doi.org/10.5281/zenodo.4895364" httpUri="https://zenodo.org/record/4895364/files/figure.png" pageId="1" pageNumber="239">Fig. 1D, E</figureCitation>
). In the older thalli these hyphae produce spores with highly distinctive multilamellate walls (
<figureCitation id="13672A40FFDCB94FB59F4F77FF39F869" box="[159,249,1896,1922]" captionStart="FIGURE 1" captionStartId="2.[151,259,1813,1837]" captionTargetBox="[155,1432,349,1790]" captionTargetId="figure-174@2.[155,1433,349,1790]" captionTargetPageId="2" captionText="FIGURE 1. The highly differentiated endophyte in Treubia pygmaea R.M.Schust.. A, B, D. Scanning electron micrographs. C, E, F. Transmission electron micrographs. A. Profile of thallus showing the intercellular fungal zone (ef) and the intracellular zone below it (if). B, C. Details of the intracellular fungus showing hyphal coils (h), fungal lumps (l) and a degenerating lump (dl). D, E. Intracellular fungus showing pseudoparenchymatous hyphae (arrowed). F. Fungal spores (s) with multilayered walls in the intercellular mucilage (*). Scale bars = (A) 200 µm; (B, D) 20 µm; (C, E, F) 5 µm." figureDoi="http://doi.org/10.5281/zenodo.4895364" httpUri="https://zenodo.org/record/4895364/files/figure.png" pageId="1" pageNumber="239">Fig. 1F</figureCitation>
). The intracellular fungal zone lies in the lower part of the thalli and comprises coils of fine hyphae some of which terminate in short-lived fungal lumps (
<figureCitation id="13672A40FFDCB94FB6304F90FC76F842" box="[816,950,1935,1961]" captionStart="FIGURE 1" captionStartId="2.[151,259,1813,1837]" captionTargetBox="[155,1432,349,1790]" captionTargetId="figure-174@2.[155,1433,349,1790]" captionTargetPageId="2" captionText="FIGURE 1. The highly differentiated endophyte in Treubia pygmaea R.M.Schust.. A, B, D. Scanning electron micrographs. C, E, F. Transmission electron micrographs. A. Profile of thallus showing the intercellular fungal zone (ef) and the intracellular zone below it (if). B, C. Details of the intracellular fungus showing hyphal coils (h), fungal lumps (l) and a degenerating lump (dl). D, E. Intracellular fungus showing pseudoparenchymatous hyphae (arrowed). F. Fungal spores (s) with multilayered walls in the intercellular mucilage (*). Scale bars = (A) 200 µm; (B, D) 20 µm; (C, E, F) 5 µm." figureDoi="http://doi.org/10.5281/zenodo.4895364" httpUri="https://zenodo.org/record/4895364/files/figure.png" pageId="1" pageNumber="239">Fig. 1B, C</figureCitation>
). Exactly the same kind of intracellular cytology is found in the epidermal and sometimes the subepidermal cells in the subterranean mucilageinvested roots of
<taxonomicName id="4C5C4D46FFDFB94CB4794887FDE0FF5A" box="[377,544,152,177]" class="Haplomitriopsida" family="Haplomitriaceae" genus="Haplomitrium" kingdom="Plantae" order="Calobryales" pageId="2" pageNumber="240" phylum="Bryophyta" rank="genus">
<emphasis id="B928EAD7FFDFB94CB4794887FDE0FF5A" box="[377,544,152,177]" italics="true" pageId="2" pageNumber="240">Haplomitrium</emphasis>
</taxonomicName>
. Spores with multilamellate walls like those in
<taxonomicName id="4C5C4D46FFDFB94CB1594887FB73FF5A" box="[1113,1203,152,177]" class="Haplomitriopsida" family="Treubiaceae" genus="Treubia" kingdom="Plantae" order="Treubiales" pageId="2" pageNumber="240" phylum="Bryophyta" rank="genus">
<emphasis id="B928EAD7FFDFB94CB1594887FB73FF5A" box="[1113,1203,152,177]" italics="true" pageId="2" pageNumber="240">Treubia</emphasis>
</taxonomicName>
can also be seen in this mucilage. Unlike all other bryophyte-fungus associations where hyphal entry is via the rhizoids, in
<taxonomicName id="4C5C4D46FFDFB94CB59748FAFF31FF15" box="[151,241,229,254]" class="Haplomitriopsida" family="Treubiaceae" genus="Treubia" kingdom="Plantae" order="Treubiales" pageId="2" pageNumber="240" phylum="Bryophyta" rank="genus">
<emphasis id="B928EAD7FFDFB94CB59748FAFF31FF15" box="[151,241,229,254]" italics="true" pageId="2" pageNumber="240">Treubia</emphasis>
</taxonomicName>
the rhizoids remain uninfected and entry is via mucilage-filled spaces between the epidermal cells. In
<taxonomicName id="4C5C4D46FFDFB94CB5974913FEFDFECE" box="[151,317,268,293]" class="Haplomitriopsida" family="Haplomitriaceae" genus="Haplomitrium" kingdom="Plantae" order="Calobryales" pageId="2" pageNumber="240" phylum="Bryophyta" rank="genus">
<emphasis id="B928EAD7FFDFB94CB5974913FEFDFECE" box="[151,317,268,293]" italics="true" pageId="2" pageNumber="240">Haplomitrium</emphasis>
</taxonomicName>
, which lacks rhizoids, hyphae penetrate directly through the epidermal cell walls.
</paragraph>
<caption id="DF23664DFFDFB94CB5974F0AFF34F808" ID-DOI="http://doi.org/10.5281/zenodo.4895364" ID-Zenodo-Dep="4895364" httpUri="https://zenodo.org/record/4895364/files/figure.png" pageId="2" pageNumber="240" startId="2.[151,259,1813,1837]" targetBox="[155,1432,349,1790]" targetPageId="2">
<paragraph id="8BE336C5FFDFB94CB5974F0AFF34F808" blockId="2.[151,1436,1813,2019]" pageId="2" pageNumber="240">
<emphasis id="B928EAD7FFDFB94CB5974F0AFED9F8C6" bold="true" box="[151,281,1813,1837]" pageId="2" pageNumber="240">FIGURE 1</emphasis>
. The highly differentiated endophyte in
<taxonomicName id="4C5C4D46FFDFB94CB7F04F09FB87F8C6" authority="R. M. Schust" authorityName="R. M. Schust" box="[752,1095,1814,1837]" class="Haplomitriopsida" family="Treubiaceae" genus="Treubia" kingdom="Plantae" order="Treubiales" pageId="2" pageNumber="240" phylum="Bryophyta" rank="species" species="pygmaea">
<emphasis id="B928EAD7FFDFB94CB7F04F09FC75F8C6" box="[752,949,1814,1837]" italics="true" pageId="2" pageNumber="240">Treubia pygmaea</emphasis>
R.M.Schust
</taxonomicName>
..
<emphasis id="B928EAD7FFDFB94CB1644F0AFB04F8C6" bold="true" box="[1124,1220,1813,1837]" pageId="2" pageNumber="240">A, B, D.</emphasis>
Scanning electron micrographs.
<emphasis id="B928EAD7FFDFB94CB4334F25FE49F8B9" bold="true" box="[307,393,1850,1874]" pageId="2" pageNumber="240">C, E, F.</emphasis>
Transmission electron micrographs.
<emphasis id="B928EAD7FFDFB94CB6294F25FC83F8B9" bold="true" box="[809,835,1850,1874]" pageId="2" pageNumber="240">A.</emphasis>
Profile of thallus showing the intercellular fungal zone (ef) and the intracellular zone below it (if).
<emphasis id="B928EAD7FFDFB94CB77A4F42FD74F89E" bold="true" box="[634,692,1885,1909]" pageId="2" pageNumber="240">B, C.</emphasis>
Details of the intracellular fungus showing hyphal coils (h), fungal lumps (l) and a degenerating lump (dl).
<emphasis id="B928EAD7FFDFB94CB7424F9DFDBAF871" bold="true" box="[578,634,1922,1946]" pageId="2" pageNumber="240">D, E.</emphasis>
Intracellular fungus showing pseudoparenchymatous hyphae (arrowed).
<emphasis id="B928EAD7FFDFB94CB0874F9DFA5CF871" bold="true" box="[1415,1436,1922,1946]" pageId="2" pageNumber="240">F.</emphasis>
Fungal spores (s) with multilayered walls in the intercellular mucilage (*). Scale bars = (A) 200 µm; (B, D) 20 µm; (C, E, F) 5 µm.
</paragraph>
</caption>
<paragraph id="8BE336C5FFDEB94DB5C64887FEFEFD69" blockId="3.[151,1437,152,642]" pageId="3" pageNumber="241">
The presence of an intercellular phase for the infections in
<taxonomicName id="4C5C4D46FFDEB94DB6A64887FBC0FF5A" box="[934,1024,152,177]" class="Haplomitriopsida" family="Treubiaceae" genus="Treubia" kingdom="Plantae" order="Treubiales" pageId="3" pageNumber="241" phylum="Bryophyta" rank="genus">
<emphasis id="B928EAD7FFDEB94DB6A64887FBC0FF5A" box="[934,1024,152,177]" italics="true" pageId="3" pageNumber="241">Treubia</emphasis>
</taxonomicName>
draws parallels with the same in
<taxonomicName id="4C5C4D46FFDEB94DB59748A0FEE8FF33" box="[151,296,191,216]" class="Lycopodiopsida" family="Lycopodiaceae" genus="Lycopodium" kingdom="Plantae" order="Lycopodiales" pageId="3" pageNumber="241" phylum="Tracheophyta" rank="genus">
<emphasis id="B928EAD7FFDEB94DB59748A0FEE8FF33" box="[151,296,191,216]" italics="true" pageId="3" pageNumber="241">Lycopodium</emphasis>
</taxonomicName>
L. (
<bibRefCitation id="EFCD4B34FFDEB94DB45948A1FD42FF33" author="Duckett, J. G. &amp; Ligrone, R." box="[345,642,190,216]" pageId="3" pageNumber="241" pagination="58 - 72" refId="ref7454" refString="Duckett, J. G. &amp; Ligrone, R. (1992) A light and electron microscope study of the fungal endophytes in the sporophyte and gametophyte of Lycopodium cernuum L. with observations on the gametophyte-sporophyte junction. Canadian Journal of Botany 70: 58 - 72." type="journal article" year="1992">Duckett &amp; Ligrone 1992</bibRefCitation>
) and even more strikingly with the Devonian plant
<taxonomicName id="4C5C4D46FFDEB94DB1EB48A0FE8AFF14" authority="Lyon ex Hoeg." authorityName="Hoeg." class="Zosterophyllopsida" family="Zosterophyllaceae" genus="Nothia" higherTaxonomySource="GBIF" kingdom="Plantae" order="Zosterophyllales" pageId="3" pageNumber="241" phylum="Tracheophyta" rank="species" species="aphylla">
<emphasis id="B928EAD7FFDEB94DB1EB48A0FA5CFF33" box="[1259,1436,191,216]" italics="true" pageId="3" pageNumber="241">Nothia aphylla</emphasis>
Lyon ex Høeg.
</taxonomicName>
(
<bibRefCitation id="EFCD4B34FFDEB94DB45B48FAFD89FF14" author="Krings, M. &amp; Taylor, T. N. &amp; Hass, H. &amp; Kerp, H. &amp; Dotzler, N. &amp; Hermsen, E. J." box="[347,585,229,255]" pageId="3" pageNumber="241" pagination="648 - 657" refId="ref8957" refString="Krings, M., Taylor, T. N., Hass, H., Kerp, H., Dotzler, N. &amp; Hermsen, E. J. (2007 a) Fungal endophytes in a 400 - millionyr-old land plant: infection pathways, spatial distribution, and host responses. New Phytologist 174: 648 - 657." type="journal article" year="2007">
Krings
<emphasis id="B928EAD7FFDEB94DB4B748FAFE34FF15" box="[439,500,229,254]" italics="true" pageId="3" pageNumber="241">et al.</emphasis>
2007a
</bibRefCitation>
, b). Given this distinctive ultrastructure and the fact that
<taxonomicName id="4C5C4D46FFDEB94DB00C48FAFAA8FF15" box="[1292,1384,229,254]" class="Haplomitriopsida" family="Treubiaceae" genus="Treubia" kingdom="Plantae" order="Treubiales" pageId="3" pageNumber="241" phylum="Bryophyta" rank="genus">
<emphasis id="B928EAD7FFDEB94DB00C48FAFAA8FF15" box="[1292,1384,229,254]" italics="true" pageId="3" pageNumber="241">Treubia</emphasis>
</taxonomicName>
and
<taxonomicName id="4C5C4D46FFDEB94DB5974913FEFDFECE" box="[151,317,268,293]" class="Haplomitriopsida" family="Haplomitriaceae" genus="Haplomitrium" kingdom="Plantae" order="Calobryales" pageId="3" pageNumber="241" phylum="Bryophyta" rank="genus">
<emphasis id="B928EAD7FFDEB94DB5974913FEFDFECE" box="[151,317,268,293]" italics="true" pageId="3" pageNumber="241">Haplomitrium</emphasis>
</taxonomicName>
are now considered to have had a very long evolutionary history in the land plant tree of life, it is extremely disappointing that the only published molecular study to date (
<bibRefCitation id="EFCD4B34FFDEB94DB115492DFB3BFEA7" author="Ligrone, R. &amp; Carafa, A. &amp; Lumini, E. &amp; Bianciotto, V. &amp; Bonfante, P. &amp; Duckett, J. G." box="[1045,1275,306,332]" pageId="3" pageNumber="241" pagination="1756 - 1777" refId="ref9253" refString="Ligrone, R., Carafa, A., Lumini, E., Bianciotto, V., Bonfante, P. &amp; Duckett, J. G. (2007) Glomeromycotean associations in liverworts: a molecular, cellular and taxonomic analysis. American Journal of Botany 94: 1756 - 1777." type="journal article" year="2007">
Ligrone
<emphasis id="B928EAD7FFDEB94DB17C492DFB77FEA0" box="[1148,1207,306,331]" italics="true" pageId="3" pageNumber="241">et al.</emphasis>
2007
</bibRefCitation>
) revealed the presence of a
<taxonomicName id="4C5C4D46FFDEB94DB4434946FE61FE99" box="[323,417,345,370]" class="Glomeromycetes" family="Glomeraceae" genus="Glomus" kingdom="Animalia" order="Glomerales" pageId="3" pageNumber="241" phylum="Glomeromycota" rank="genus">
<emphasis id="B928EAD7FFDEB94DB4434946FE61FE99" box="[323,417,345,370]" italics="true" pageId="3" pageNumber="241">Glomus</emphasis>
</taxonomicName>
in
<taxonomicName id="4C5C4D46FFDEB94DB4CA4946FC4BFE98" authority="R. M. Schust." authorityName="R. M. Schust." box="[458,907,345,371]" class="Haplomitriopsida" family="Haplomitriaceae" genus="Haplomitrium" kingdom="Plantae" order="Calobryales" pageId="3" pageNumber="241" phylum="Bryophyta" rank="species" species="chilensis">
<emphasis id="B928EAD7FFDEB94DB4CA4946FD2AFE99" box="[458,746,345,370]" italics="true" pageId="3" pageNumber="241">Haplomitrium chilensis</emphasis>
R.M.Schust.
</taxonomicName>
closely allied to the
<taxonomicName id="4C5C4D46FFDEB94DB18E4946FA5DFE98" authority="Tul. &amp; C. Tul." authorityName="Tul. &amp; C. Tul." box="[1166,1437,345,371]" class="Glomeromycetes" family="Glomeraceae" genus="Glomus" kingdom="Animalia" order="Glomerales" pageId="3" pageNumber="241" phylum="Glomeromycota" rank="genus">
<emphasis id="B928EAD7FFDEB94DB18E4946FB2CFE99" box="[1166,1260,345,370]" italics="true" pageId="3" pageNumber="241">Glomus</emphasis>
Tul. &amp; C.Tul.
</taxonomicName>
Group A endophytes found in
<taxonomicName id="4C5C4D46FFDEB94DB4FF499FFD25FE71" authority="Hill" authorityName="Hill" box="[511,741,384,410]" class="Marchantiopsida" family="Conocephalaceae" genus="Conocephalum" kingdom="Plantae" order="Marchantiales" pageId="3" pageNumber="241" phylum="Bryophyta" rank="genus">
<emphasis id="B928EAD7FFDEB94DB4FF499FFD70FE72" box="[511,688,384,409]" italics="true" pageId="3" pageNumber="241">Conocephalum</emphasis>
Hill
</taxonomicName>
,
<taxonomicName id="4C5C4D46FFDEB94DB7F5499FFC2FFE71" authority="Hazsl." authorityName="Hazsl." box="[757,1007,384,410]" class="Jungermanniopsida" family="Fossombroniaceae" genus="Fossombronia" kingdom="Plantae" order="Fossombroniales" pageId="3" pageNumber="241" phylum="Bryophyta" rank="genus">
<emphasis id="B928EAD7FFDEB94DB7F5499FFC5DFE72" box="[757,925,384,409]" italics="true" pageId="3" pageNumber="241">Fossombronia</emphasis>
Hazsl.
</taxonomicName>
and
<taxonomicName id="4C5C4D46FFDEB94DB129499FFAF9FE71" authority="Raddi. However" authorityName="Raddi. However" box="[1065,1337,384,410]" class="Jungermanniopsida" family="Pelliaceae" genus="Pellia" kingdom="Plantae" order="Pelliales" pageId="3" pageNumber="241" phylum="Bryophyta" rank="genus">
<emphasis id="B928EAD7FFDEB94DB129499FFBAFFE72" box="[1065,1135,384,409]" italics="true" pageId="3" pageNumber="241">Pellia</emphasis>
Raddi. However
</taxonomicName>
, a much more extensive sequencing programme of the
<taxonomicName id="4C5C4D46FFDEB94DB7E149B8FC81FE2B" box="[737,833,423,448]" class="Haplomitriopsida" family="Treubiaceae" genus="Treubia" kingdom="Plantae" order="Treubiales" pageId="3" pageNumber="241" phylum="Bryophyta" rank="genus">
<emphasis id="B928EAD7FFDEB94DB7E149B8FC81FE2B" box="[737,833,423,448]" italics="true" pageId="3" pageNumber="241">Treubia</emphasis>
</taxonomicName>
and
<taxonomicName id="4C5C4D46FFDEB94DB68049B8FBF0FE2B" box="[896,1072,423,448]" class="Haplomitriopsida" family="Haplomitriaceae" genus="Haplomitrium" kingdom="Plantae" order="Calobryales" pageId="3" pageNumber="241" phylum="Bryophyta" rank="genus">
<emphasis id="B928EAD7FFDEB94DB68049B8FBF0FE2B" box="[896,1072,423,448]" italics="true" pageId="3" pageNumber="241">Haplomitrium</emphasis>
</taxonomicName>
endophytes now in progress (
<bibRefCitation id="EFCD4B34FFDEB94DB59F49D2FE2BFE0C" author="Bidartondo, M. I. &amp; Duckett, J. G." box="[159,491,461,487]" pageId="3" pageNumber="241" pagination="485 - 492" refId="ref7007" refString="Bidartondo, M. I. &amp; Duckett, J. G. (2010) Conservative ecological and evolutionary patterns in liverwort-fungal symbioses. Proceedings of the Royal Society B. 277: 485 - 492." type="journal article" year="2010">Bidartondo &amp; Duckett 2010</bibRefCitation>
, unpublished data) is revealing that these two genera, plus several primitive taxa in the thalloid liverworts, lack
<taxonomicName id="4C5C4D46FFDEB94DB71449EBFDB2FDE6" box="[532,626,500,525]" class="Glomeromycetes" family="Glomeraceae" genus="Glomus" kingdom="Animalia" order="Glomerales" pageId="3" pageNumber="241" phylum="Glomeromycota" rank="genus">
<emphasis id="B928EAD7FFDEB94DB71449EBFDB2FDE6" box="[532,626,500,525]" italics="true" pageId="3" pageNumber="241">Glomus</emphasis>
</taxonomicName>
but contain zygomycetous endophytes, a group of fungi recently resolved as ancestral to the
<taxonomicName id="4C5C4D46FFDEB94DB4D74A05FD59FDDF" box="[471,665,538,564]" class="Glomeromycetes" kingdom="Animalia" pageId="3" pageNumber="238" phylum="Glomeromycota" rank="class">glomeromycetes</taxonomicName>
(
<bibRefCitation id="EFCD4B34FFDEB94DB7A84A05FCB8FDDF" author="James, T. Y. &amp; Kauff, F. &amp; Schoch, C. L. &amp; Matheny, P. B. &amp; Hofstetter, V. &amp; Cox, C. J. &amp; Celio, G. &amp; Gueidan, C. &amp; Fraker, E. &amp; Miadlikowska, J. &amp; Lumbsch, H. T. &amp; Rauhut, A. &amp; Reeb, V. &amp; Arnold, A. E. &amp; Amtoft, A. &amp; Stajich, J. E. &amp; Hosakam, K. &amp; Sung, G-H. &amp; Johnson, D. &amp; O'Rourke, B. &amp; Crockett, M. &amp; Binder, M. &amp; Curtis, J. M. &amp; Slot, J. C. &amp; Wang, Z. &amp; Wilson, A. W. &amp; Schussler, A. &amp; Longcore, J. E. &amp; O'Donnell, K. &amp; Mozley-Standridge, S. &amp; Porter, D. &amp; Letcher, P. M. &amp; Powell, M. J. &amp; Taylor, J. W. &amp; White, M. M. &amp; Griffith, G. W. &amp; Davies, D. R. &amp; Humber, R. A. &amp; Morton, J. B. &amp; Sugiyama, J. &amp; Rossman, A. Y. &amp; Rogers, J. D. &amp; Pfister, D. H. &amp; Hewitt, D. &amp; Hansen, K. &amp; Hambleton, S. &amp; Shoemaker, R. A. &amp; Kohlmeyer, J. &amp; Volkmann-Kohlmeyer, B. &amp; Spotts, R. &amp; Serdani, M. &amp; Crous, P. W. &amp; Hughes, K. W. &amp; Matsuura, K. &amp; Langer, E. &amp; Langer, G. &amp; Untereiner, W. A. &amp; Lucking, R. &amp; Budel, B. &amp; Geiser, D. M. &amp; Aptroot, A. &amp; Diederich, P. &amp; Schmitt, I. &amp; Schultz, M. &amp; Yahr, R. &amp; Hibbett, D. S. &amp; Lutzoni, F. &amp; McLaughlin, D. J. &amp; Spatafora, J. W. &amp; Vilgalys, R." box="[680,888,538,564]" pageId="3" pageNumber="241" pagination="818 - 822" refId="ref8432" refString="James, T. Y., Kauff, F., Schoch, C. L., Matheny, P. B., Hofstetter, V., Cox, C. J., Celio, G., Gueidan, C., Fraker, E., Miadlikowska, J., Lumbsch, H. T., Rauhut, A., Reeb, V., Arnold, A. E., Amtoft, A., Stajich, J. E., Hosakam, K., Sung, G-H., Johnson, D., O'Rourke, B., Crockett, M., Binder, M., Curtis, J. M., Slot, J. C., Wang, Z., Wilson, A. W., Schussler, A., Longcore, J. E., O'Donnell, K., Mozley-Standridge, S., Porter, D., Letcher, P. M., Powell, M. J., Taylor, J. W., White, M. M., Griffith, G. W., Davies, D. R., Humber, R. A., Morton, J. B., Sugiyama, J., Rossman, A. Y., Rogers, J. D., Pfister, D. H., Hewitt, D., Hansen, K., Hambleton, S., Shoemaker, R. A., Kohlmeyer, J., Volkmann-Kohlmeyer, B., Spotts, R., Serdani, M., Crous, P. W., Hughes, K. W., Matsuura, K., Langer, E., Langer, G., Untereiner, W. A., Lucking, R., Budel, B., Geiser, D. M., Aptroot, A., Diederich, P., Schmitt, I., Schultz, M., Yahr, R., Hibbett, D. S., Lutzoni, F., McLaughlin, D. J., Spatafora, J. W. &amp; Vilgalys, R. (2006) Reconstructing the early evolution of Fungi using a six-gene phylogeny. Nature 443: 818 - 822." type="journal article" year="2006">
James
<emphasis id="B928EAD7FFDEB94DB7F84A05FCF2FDD8" box="[760,818,538,563]" italics="true" pageId="3" pageNumber="241">et al.</emphasis>
2006
</bibRefCitation>
). Thus, we now have the first glimpses of what may well turn out to be a hitherto unsuspected spectrum of novel fungal symbioses very early in the ancestry of land plants.
</paragraph>
</subSubSection>
</treatment>
</document>