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<document id="911CAC75740159B47D84992D2E0E62AE" ID-DOI="10.11646/phytotaxa.664.2.9" ID-ISSN="1179-3163" ID-Zenodo-Dep="14517438" IM.bibliography_approvedBy="felipe" IM.materialsCitations_approvedBy="guilherme" IM.metadata_approvedBy="guilherme" IM.taxonomicNames_approvedBy="guilherme" IM.treatments_approvedBy="guilherme" checkinTime="1734542953762" checkinUser="felipe" docAuthor="Hu, Yu-Ping, Zhang, Li-Bing &amp; Zhou, Xin-Mao" docDate="2024" docId="03BF87A8AF54FFC0FF0454899B75063F" docLanguage="en" docName="phytotaxa.664.2.155-158.pdf" docOrigin="Phytotaxa 664 (2)" docSource="https://doi.org/10.11646/phytotaxa.664.2.9" docStyle="DocumentStyle:96748F8F1B6C902996E134952A3A36B9.13:Phytotaxa.2014-.journal_article" docStyleId="96748F8F1B6C902996E134952A3A36B9" docStyleName="Phytotaxa.2014-.journal_article" docStyleVersion="13" docTitle="Didiclis khokhrjakovii Li Bing Zhang &amp; X. M. Zhou 2024, comb. nov." docType="treatment" docVersion="3" lastPageNumber="156" masterDocId="FF86FFD0AF55FFC1FF8C54169F1B0420" masterDocTitle="Asian Selaginellaceae: Five new combinations in Didiclis and Hypopterygiopsis" masterLastPageNumber="158" masterPageNumber="155" pageNumber="156" updateTime="1740768937979" updateUser="ExternalLinkService" zenodo-license-figures="UNSPECIFIED" zenodo-license-treatments="UNSPECIFIED">
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<mods:title id="0D4A997213ABB9DD007CBB610E5A8FAF">Asian Selaginellaceae: Five new combinations in Didiclis and Hypopterygiopsis</mods:title>
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<mods:namePart id="D59B9C778B55EE53D499FA556DCBCAD5">Hu, Yu-Ping</mods:namePart>
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<mods:namePart id="E0AD0BFD1E9A5218E73D03764EF354D3">Zhang, Li-Bing</mods:namePart>
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<mods:affiliation id="98EBFE16B5469B3F3F8BA3CF91EF87EE">Missouri Botanical Garden, 4344 Shaw Blvd, St. Louis, Missouri 63110, U. S. A.; Chengdu Institute of Biology, Chinese Academy of Sciences, P. O. Box 416, Chengdu, Sichuan 610041, China</mods:affiliation>
<mods:nameIdentifier id="B63A88350368391BA9FF2CC3267D84F2" type="email">Libing.Zhang@mobot.org</mods:nameIdentifier>
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<mods:namePart id="2E317F7D186C0FE21A61947CB270AFBC">Zhou, Xin-Mao</mods:namePart>
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<taxonomicName id="4C164D3DAF54FFC0FF0454899C2E0497" authority="(Shalimov) Li Bing Zhang &amp; X. M. Zhou" authorityName="Li Bing Zhang &amp; X. M. Zhou" authorityYear="2024" baseAuthorityName="Shalimov" box="[136,821,159,183]" class="Lycopodiopsida" family="Selaginellaceae" genus="Didiclis" kingdom="Plantae" order="Selaginellales" pageId="1" pageNumber="156" phylum="Tracheophyta" rank="species" species="khokhrjakovii" status="comb. nov.">
<emphasis id="B962EAACAF54FFC0FF0454899E670497" bold="true" box="[136,380,159,183]" italics="true" pageId="1" pageNumber="156">Didiclis khokhrjakovii</emphasis>
(Shalimov) Li Bing Zhang &amp; X.M. Zhou
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,
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<paragraph id="8BA936BEAF54FFC0FC3A54899EDC04FB" blockId="1.[136,1451,159,255]" pageId="1" pageNumber="156">
<collectingMunicipality id="6BCDACC4AF54FFC0FC3A54899B3D0497" box="[950,1062,159,183]" pageId="1" pageNumber="156">Basionym</collectingMunicipality>
:
<taxonomicName id="4C164D3DAF54FFC0FBA354899AB00497" authority="Shalimov" authorityName="Shalimov" box="[1071,1451,159,183]" class="Lycopodiopsida" family="Selaginellaceae" genus="Selaginella" kingdom="Plantae" order="Selaginellales" pageId="1" pageNumber="156" phylum="Tracheophyta" rank="species" species="khokhrjakovii">
<emphasis id="B962EAACAF54FFC0FBA354899A5B0497" box="[1071,1344,159,183]" italics="true" pageId="1" pageNumber="156">Selaginella khokhrjakovii</emphasis>
Shalimov
</taxonomicName>
in
<bibRefCitation id="EF874B4FAF54FFC0FF2854D59ED804FB" author="Shalimov, A. P. &amp; Zaikov, V. F. &amp; Kalyuzhny, S. S. &amp; Zhang, M. - H. &amp; Zhang, X. - C." box="[164,451,195,219]" pageId="1" pageNumber="156" pagination="45 - 54" refId="ref2075" refString="Shalimov, A. P., Zaikov, V. F., Kalyuzhny, S. S., Zhang, M. - H. &amp; Zhang, X. - C. (2024) The recovery of Selaginella flaccida based on morphological and molecular data, with the description of S. khokhrjakovii from Indo-China and Peninsular Malaysia. Botanica Pacifica 13: 45 - 54. https: // doi. org / 10.17581 / bp. 2024.13105" type="journal article" year="2024">
Shalimov
<emphasis id="B962EAACAF54FFC0FE9E54D29E5304FB" box="[274,328,195,219]" italics="true" pageId="1" pageNumber="156">et al.</emphasis>
(2024: 51)
</bibRefCitation>
.
</paragraph>
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<subSubSection id="C30C6535AF54FFC0FE4154D29B4504DF" pageId="1" pageNumber="156" type="materials_examined">
<paragraph id="8BA936BEAF54FFC0FE4154D29B4504DF" blockId="1.[136,1451,159,255]" pageId="1" pageNumber="156">
<location id="8EC96065AF54FFC0FE4154D29D1D04FC" LSID="urn:lsid:plazi:treatment:03BF87A8AF54FFC0FF0454899B75063F:8EC96065AF54FFC0FE4154D29D1D04FC" box="[461,518,196,220]" country="Vietnam" municipality="Basionym" name="Type" pageId="1" pageNumber="156" stateProvince="Ho Chi Minh City">Type</location>
:
<collectingCountry id="F301762EAF54FFC0FD9C54D59D9404FB" box="[528,655,195,219]" name="Vietnam" pageId="1" pageNumber="156">VIETNAM</collectingCountry>
:
<collectingRegion id="49D2F85CAF54FFC0FD1654D59C4404FB" box="[666,863,195,219]" country="Vietnam" name="Ho Chi Minh City" pageId="1" pageNumber="156">Hô Chí Minh City</collectingRegion>
(
<collectingRegion id="49D2F85CAF54FFC0FCE254D59CA504FB" box="[878,958,195,219]" country="Vietnam" name="Ho Chi Minh City" pageId="1" pageNumber="156">Saigon</collectingRegion>
): plantations, zoo territory,
<date id="FFA8107EAF54FFC0FB6A54D59ABC04FB" box="[1254,1447,195,219]" pageId="1" pageNumber="156" value="1979-12-03">
<collectingDate id="EFECE996AF54FFC0FB6A54D59ABC04FB" box="[1254,1447,195,219]" pageId="1" pageNumber="156" value="1979-12-03">3 December 1979</collectingDate>
</date>
,
<emphasis id="B962EAACAF54FFC0FF0454F19E6804DF" box="[136,371,231,255]" italics="true" pageId="1" pageNumber="156">
<collectorName id="26E35368AF54FFC0FF0454F19E5C04DF" box="[136,327,231,255]" pageId="1" pageNumber="156">A.P. Khokhryakov</collectorName>
s.n.
</emphasis>
(
<typeStatus id="54AD881CAF54FFC0FE0D54F19EFB04DF" box="[385,480,231,255]" pageId="1" pageNumber="156" type="holotype">holotype</typeStatus>
MHA0034906!;
<typeStatus id="54AD881CAF54FFC0FD1554F19DEB04DF" box="[665,752,231,255]" pageId="1" pageNumber="156" type="isotype">isotypes</typeStatus>
MHA0034907!, MHA0034908!).
</paragraph>
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<paragraph id="8BA936BEAF54FFC0FF0455399BC30567" blockId="1.[136,1452,303,543]" box="[136,1240,303,328]" pageId="1" pageNumber="156">
<emphasis id="B962EAACAF54FFC0FF0455399E010567" bold="true" box="[136,282,303,327]" inLineHeading="true" pageId="1" pageNumber="156" reason="1">Distribution.</emphasis>
<collectingCountry id="F301762EAF54FFC0FEAD55399E8A0567" box="[289,401,303,327]" name="Cambodia" pageId="1" pageNumber="156">Cambodia</collectingCountry>
, (?)
<collectingCountry id="F301762EAF54FFC0FE4C55399EE20567" box="[448,505,303,327]" name="India" pageId="1" pageNumber="156">India</collectingCountry>
,
<collectingCountry id="F301762EAF54FFC0FD8855399D730567" box="[516,616,303,327]" name="Malaysia" pageId="1" pageNumber="156">Malaysia</collectingCountry>
,
<collectingCountry id="F301762EAF54FFC0FDFF55269DC70568" box="[627,732,304,328]" name="Myanmar" pageId="1" pageNumber="156">Myanmar</collectingCountry>
,
<collectingCountry id="F301762EAF54FFC0FD6B55399C520567" box="[743,841,303,327]" name="Thailand" pageId="1" pageNumber="156">Thailand</collectingCountry>
, and
<collectingCountry id="F301762EAF54FFC0FCF355399CC20567" box="[895,985,303,327]" name="Vietnam" pageId="1" pageNumber="156">Vietnam</collectingCountry>
(
<bibRefCitation id="EF874B4FAF54FFC0FC6B55399BD70567" author="Shalimov, A. P. &amp; Zaikov, V. F. &amp; Kalyuzhny, S. S. &amp; Zhang, M. - H. &amp; Zhang, X. - C." box="[999,1228,303,327]" pageId="1" pageNumber="156" pagination="45 - 54" refId="ref2075" refString="Shalimov, A. P., Zaikov, V. F., Kalyuzhny, S. S., Zhang, M. - H. &amp; Zhang, X. - C. (2024) The recovery of Selaginella flaccida based on morphological and molecular data, with the description of S. khokhrjakovii from Indo-China and Peninsular Malaysia. Botanica Pacifica 13: 45 - 54. https: // doi. org / 10.17581 / bp. 2024.13105" type="journal article" year="2024">
Shalimov
<emphasis id="B962EAACAF54FFC0FBDA55269B970567" box="[1110,1164,303,327]" italics="true" pageId="1" pageNumber="156">et al.</emphasis>
2024
</bibRefCitation>
).
</paragraph>
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<subSubSection id="C30C6535AF54FFC0FF3155459B75063F" pageId="1" pageNumber="156" type="discussion">
<paragraph id="8BA936BEAF54FFC0FF3155459B75063F" blockId="1.[136,1452,303,543]" pageId="1" pageNumber="156">
<emphasis id="B962EAACAF54FFC0FF3155459FE6054B" bold="true" box="[189,253,339,363]" pageId="1" pageNumber="156">Notes</emphasis>
: This species is similar to
<taxonomicName id="4C164D3DAF54FFC0FDA455459DC7054B" authorityName="Li Bing Zhang &amp; X. M. Zhou" authorityYear="2024" baseAuthorityName="Li Bing Zhang &amp; X. M. Zhou" baseAuthorityYear="2024" box="[552,732,339,363]" class="Lycopodiopsida" family="Selaginellaceae" genus="Didiclis" kingdom="Plantae" order="Selaginellales" pageId="1" pageNumber="156" phylum="Tracheophyta" rank="species" species="flaccida">
<emphasis id="B962EAACAF54FFC0FDA455459DC7054B" box="[552,732,339,363]" italics="true" pageId="1" pageNumber="156">Didiclis flaccida</emphasis>
</taxonomicName>
in having main stems thick, primary leafy branches
<quantity id="4CEE9B5BAF54FFC0FAAE55459A6C054B" box="[1314,1399,339,363]" metricMagnitude="-2" metricUnit="m" metricValue="5.0" metricValueMax="8.0" metricValueMin="2.0" pageId="1" pageNumber="156" unit="cm" value="5.0" valueMax="8.0" valueMin="2.0">28 cm</quantity>
, lax, secondary branches simple or rarely forked; dorsal leaves oblique, elliptic, narrowly ovate-elliptic or falcate, margins entire to subentire, and apex acuminate to cuspidate. However,
<taxonomicName id="4C164D3DAF54FFC0FC88558A9CA00593" box="[772,955,411,435]" class="Lycopodiopsida" family="Selaginellaceae" genus="Selaginella" kingdom="Plantae" order="Selaginellales" pageId="1" pageNumber="156" phylum="Tracheophyta" rank="species" species="khokhrjakovii">
<emphasis id="B962EAACAF54FFC0FC88558A9CA00593" box="[772,955,411,435]" italics="true" pageId="1" pageNumber="156">D. khokhrjakovii</emphasis>
</taxonomicName>
has ventral leaves ovate, acuminate at apex, margins entire to subentire (with spaced teeth); dorsal leaves entire to subentire (with spaced toothed), apex acuminate. In contrast,
<taxonomicName id="4C164D3DAF54FFC0FF6655F29E7B05DB" authorityName="Li Bing Zhang &amp; X. M. Zhou" authorityYear="2024" baseAuthorityName="Li Bing Zhang &amp; X. M. Zhou" baseAuthorityYear="2024" box="[234,352,483,507]" class="Lycopodiopsida" family="Selaginellaceae" genus="Didiclis" kingdom="Plantae" order="Selaginellales" pageId="1" pageNumber="156" phylum="Tracheophyta" rank="species" species="flaccida">
<emphasis id="B962EAACAF54FFC0FF6655F29E7B05DB" box="[234,352,483,507]" italics="true" pageId="1" pageNumber="156">D. flaccida</emphasis>
</taxonomicName>
has ventral leaves oblong-ovate, subacute at apex, margins entire at base, minutely denticulate towards at apex; dorsal leaves, margin entire, and apex acuminate to cuspidate (
<bibRefCitation id="EF874B4FAF54FFC0FCF256119B78063F" author="Shalimov, A. P. &amp; Zaikov, V. F. &amp; Kalyuzhny, S. S. &amp; Zhang, M. - H. &amp; Zhang, X. - C." box="[894,1123,519,543]" pageId="1" pageNumber="156" pagination="45 - 54" refId="ref2075" refString="Shalimov, A. P., Zaikov, V. F., Kalyuzhny, S. S., Zhang, M. - H. &amp; Zhang, X. - C. (2024) The recovery of Selaginella flaccida based on morphological and molecular data, with the description of S. khokhrjakovii from Indo-China and Peninsular Malaysia. Botanica Pacifica 13: 45 - 54. https: // doi. org / 10.17581 / bp. 2024.13105" type="journal article" year="2024">
Shalimov
<emphasis id="B962EAACAF54FFC0FC61561E9B38063F" box="[1005,1059,519,543]" italics="true" pageId="1" pageNumber="156">et al.</emphasis>
2024
</bibRefCitation>
).
</paragraph>
</subSubSection>
</treatment>
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<mods:namePart id="1B8E530E9B6022962588D4AF782D6721">Hu, Yu-Ping</mods:namePart>
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<mods:namePart id="82620D341ED8F9843D090860219EFED9">Zhang, Li-Bing</mods:namePart>
<mods:nameIdentifier id="92960C796E91271F4340659261978D04" type="ORCID">0000-0002-4905-040X</mods:nameIdentifier>
<mods:affiliation id="FEB823E21A1445E6A38120BDCA30440D">Missouri Botanical Garden, 4344 Shaw Blvd, St. Louis, Missouri 63110, U. S. A.; Chengdu Institute of Biology, Chinese Academy of Sciences, P. O. Box 416, Chengdu, Sichuan 610041, China</mods:affiliation>
<mods:nameIdentifier id="5F65005C5BD213CD4946A23A4697D56A" type="email">Libing.Zhang@mobot.org</mods:nameIdentifier>
</mods:name>
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<mods:namePart id="97E87AFBCCF6E75C8C447718D3455976">Zhou, Xin-Mao</mods:namePart>
<mods:nameIdentifier id="741C3F8BA72C4841EC503E8A0DC401D5" type="ORCID">0000-0003-3555-7784</mods:nameIdentifier>
<mods:affiliation id="20AE69CEAC761D08E23FE2705411BA5A">School of Ecology and Environmental Science, Yunnan University, Kunming 650504, Yunnan, China</mods:affiliation>
<mods:nameIdentifier id="B02B6E47360964EF67A04012773C0736" type="email">xinmao.zhou@ynu.edu.cn</mods:nameIdentifier>
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<mods:title id="5C179D7F4294DF5FD2EDC922BC85C0EB">Phytotaxa</mods:title>
</mods:titleInfo>
<mods:part id="9736E22A47CDC662C5245A73E1D0225D">
<mods:date id="16899FC2D2D46A840FC45CFDC169A8D5">2024</mods:date>
<mods:detail id="76431755B79C149521715D87240F0D11" type="pubDate">
<mods:number id="8AB8E376317F9BA0162DB4884167494D">2024-09-05</mods:number>
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<mods:detail id="B79CD9B316C562CA029F4551894DD1DD" type="volume">
<mods:number id="DB09B7C0FA1EB9D796F14B18B1DCF1D7">664</mods:number>
</mods:detail>
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<mods:number id="94836A6E8F4B5139A3E096B9806D344C">2</mods:number>
</mods:detail>
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<mods:start id="46FB6AA367191A6011BCA98234831245">155</mods:start>
<mods:end id="E84898C488D22451E025FC52F7A73211">158</mods:end>
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<mods:location id="DB9830ECA58F7DE217F3C5927E262855">
<mods:url id="F41D6636F7861A8FDC3AA688D525C429">https://doi.org/10.11646/phytotaxa.664.2.9</mods:url>
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<mods:identifier id="223D0C1450658FC264C6975673CCA082" type="ISSN">1179-3163</mods:identifier>
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<taxonomicName id="4C164D3DAF55FFC1FF0452A59D8102EA" authority="(Bory) Li Bing Zhang &amp; X. M. Zhou" authorityName="Li Bing Zhang &amp; X. M. Zhou" authorityYear="2024" baseAuthorityName="Bory" box="[136,666,1715,1738]" class="Lycopodiopsida" family="Selaginellaceae" genus="Didiclis" kingdom="Plantae" order="Selaginellales" pageId="0" pageNumber="155" phylum="Tracheophyta" rank="species" species="flaccida" status="comb. nov.">
<emphasis id="B962EAACAF55FFC1FF0452A59E6F02E9" bold="true" box="[136,372,1715,1737]" pageId="0" pageNumber="155">
<emphasis id="B962EAACAF55FFC1FF0452A59E3102E9" bold="true" box="[136,298,1715,1737]" italics="true" pageId="0" pageNumber="155">Didiclis flaccida</emphasis>
(Bory)
</emphasis>
Li Bing Zhang &amp; X.M. Zhou
</taxonomicName>
,
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<taxonomicNameLabel id="A25157D7AF55FFC1FD2952A29C1402E9" box="[677,783,1716,1737]" pageId="0" pageNumber="155" rank="species">comb. nov.</taxonomicNameLabel>
</emphasis>
</paragraph>
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<paragraph id="8BA936BEAF55FFC1FC9952A59A1602E9" blockId="0.[136,1452,1676,1774]" box="[789,1293,1714,1738]" pageId="0" pageNumber="155">
<emphasis id="B962EAACAF55FFC1FC9952A59C9902E9" bold="true" box="[789,898,1715,1737]" pageId="0" pageNumber="155">Basionym:</emphasis>
<taxonomicName id="4C164D3DAF55FFC1FC0552A29A1202E9" authority="Bory (1833: 9)" authorityName="Bory" authorityPageNumber="9" authorityYear="1833" box="[905,1289,1714,1738]" class="Lycopodiopsida" family="Lycopodiaceae" genus="Lycopodium" kingdom="Plantae" order="Lycopodiales" pageId="0" pageNumber="155" phylum="Tracheophyta" rank="species" species="flaccidum">
<emphasis id="B962EAACAF55FFC1FC0552A29B7E02E9" box="[905,1125,1716,1737]" italics="true" pageId="0" pageNumber="155">Lycopodium flaccidum</emphasis>
<bibRefCitation id="EF874B4FAF55FFC1FBE052A29A1202E9" author="Bory, J. B. G. M." box="[1132,1289,1714,1738]" pageId="0" pageNumber="155" pagination="1825 - 1829" refId="ref1698" refString="Bory, J. B. G. M. (1833) Lycopodiacees. In: Belanger, C. &amp; Bory, J. B. G. M. (Eds.) Voyage aux Indes-orientales, pendant les annees 1825 - 1829. A. Bertrand, Paris, pp. 7 - 13." type="book chapter" year="1833">
<emphasis id="B962EAACAF55FFC1FBE052A29B8402E9" bold="true" box="[1132,1183,1716,1737]" pageId="0" pageNumber="155">Bory</emphasis>
(1833:
<emphasis id="B962EAACAF55FFC1FB7F52A59A1202E9" bold="true" box="[1267,1289,1715,1737]" pageId="0" pageNumber="155">9)</emphasis>
</bibRefCitation>
</taxonomicName>
<emphasis id="B962EAACAF55FFC1FA8452A59A1602E9" bold="true" box="[1288,1293,1715,1737]" pageId="0" pageNumber="155">.</emphasis>
</paragraph>
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<paragraph id="8BA936BEAF55FFC1FA9952A49ABF02CE" blockId="0.[136,1452,1676,1774]" pageId="0" pageNumber="155">
Type:
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<collectingCountry id="F301762EAF55FFC1FAD552A49AB302EA" box="[1369,1448,1714,1738]" name="India" pageId="0" pageNumber="155">INDIA</collectingCountry>
.
<collectingMunicipality id="6BCDACC4AF55FFC1FF0452C09FDC02CE" box="[136,199,1750,1774]" pageId="0" pageNumber="155">Mahé</collectingMunicipality>
, sur la terre, dans les lieux ombragés et humides,
<emphasis id="B962EAACAF55FFC1FD5552C09C8202CE" box="[729,921,1750,1774]" italics="true" pageId="0" pageNumber="155">
<collectorName id="26E35368AF55FFC1FD5552C09C7502CE" box="[729,878,1750,1774]" pageId="0" pageNumber="155">C.P. Bélanger</collectorName>
s.n.
</emphasis>
(
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<specimenCode id="DBB09EC5AF55FFC1FB8152C09B9402CE" box="[1037,1167,1750,1774]" pageId="0" pageNumber="155">P00677428</specimenCode>
!
</materialsCitation>
;
<materialsCitation id="3B7E3CE3AF55FFC1FB1152C09A8202CE" box="[1181,1433,1750,1774]" collectionCode="WAG" collectorName="C. P. Belanger" country="India" location="Mahe" municipality="Mahe" pageId="0" pageNumber="155" specimenCode="WAG1901915" specimenCount="1" typeStatus="isotype">
<typeStatus id="54AD881CAF55FFC1FB1152C09BF102CE" box="[1181,1258,1750,1774]" pageId="0" pageNumber="155" type="isotype">isotype</typeStatus>
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!
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).
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<paragraph id="8BA936BEAF55FFC1FF0453089B9C0316" blockId="0.[136,1452,1822,2026]" box="[136,1159,1822,1846]" pageId="0" pageNumber="155">
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<emphasis id="B962EAACAF55FFC1FF0453089E080316" bold="true" box="[136,275,1822,1846]" pageId="0" pageNumber="155">Distribution</emphasis>
.
</emphasis>
<collectingCountry id="F301762EAF55FFC1FEAD53089E430316" box="[289,344,1822,1846]" name="India" pageId="0" pageNumber="155">India</collectingCountry>
(
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,
<collectingRegion id="49D2F85CAF55FFC1FE7653089D5F0316" box="[506,580,1822,1846]" country="India" name="Kerala" pageId="0" pageNumber="155">Kerala</collectingRegion>
,
<collectingRegion id="49D2F85CAF55FFC1FDC253089DA50316" box="[590,702,1822,1846]" country="India" name="Karnataka" pageId="0" pageNumber="155">Karnataka</collectingRegion>
,
<collectingRegion id="49D2F85CAF55FFC1FD4453089C520316" box="[712,841,1822,1846]" country="India" name="Tamil Nadu" pageId="0" pageNumber="155">Tamil Nadu</collectingRegion>
, Goa) (
<bibRefCitation id="EF874B4FAF55FFC1FC1B53089B670316" author="Shalimov, A. P. &amp; Zaikov, V. F. &amp; Kalyuzhny, S. S. &amp; Zhang, M. - H. &amp; Zhang, X. - C." box="[919,1148,1822,1846]" pageId="0" pageNumber="155" pagination="45 - 54" refId="ref2075" refString="Shalimov, A. P., Zaikov, V. F., Kalyuzhny, S. S., Zhang, M. - H. &amp; Zhang, X. - C. (2024) The recovery of Selaginella flaccida based on morphological and molecular data, with the description of S. khokhrjakovii from Indo-China and Peninsular Malaysia. Botanica Pacifica 13: 45 - 54. https: // doi. org / 10.17581 / bp. 2024.13105" type="journal article" year="2024">
Shalimov
<emphasis id="B962EAACAF55FFC1FB8A53099B270316" box="[1030,1084,1822,1846]" italics="true" pageId="0" pageNumber="155">et al.</emphasis>
2024
</bibRefCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="C30C6535AF55FFC1FF3153549DD803C9" pageId="0" pageNumber="155" type="discussion">
<paragraph id="8BA936BEAF55FFC1FF3153549DD803C9" blockId="0.[136,1452,1822,2026]" pageId="0" pageNumber="155">
<emphasis id="B962EAACAF55FFC1FF3153549FE6037A" bold="true" box="[189,253,1858,1882]" pageId="0" pageNumber="155">Notes</emphasis>
:
<taxonomicName id="4C164D3DAF55FFC1FE8253549ED9037A" authorityName="Li Bing Zhang &amp; X. M. Zhou" authorityYear="2024" baseAuthorityName="Li Bing Zhang &amp; X. M. Zhou" baseAuthorityYear="2024" box="[270,450,1858,1882]" class="Lycopodiopsida" family="Selaginellaceae" genus="Didiclis" kingdom="Plantae" order="Selaginellales" pageId="0" pageNumber="155" phylum="Tracheophyta" rank="species" species="flaccida">
<emphasis id="B962EAACAF55FFC1FE8253549ED9037A" box="[270,450,1858,1882]" italics="true" pageId="0" pageNumber="155">Didiclis flaccida</emphasis>
</taxonomicName>
has long been treated as a synonym of
<taxonomicName id="4C164D3DAF55FFC1FC0C53559B14037A" box="[896,1039,1858,1882]" class="Lycopodiopsida" family="Selaginellaceae" genus="Didiclis" kingdom="Plantae" order="Selaginellales" pageId="0" pageNumber="155" phylum="Tracheophyta" rank="species" species="delicatula">
<emphasis id="B962EAACAF55FFC1FC0C53559B14037A" box="[896,1039,1858,1882]" italics="true" pageId="0" pageNumber="155">D. delicatula</emphasis>
</taxonomicName>
(Desvaux ex Poiret in Lamark 1814: 554) Li Bing Zhang &amp; X.M. Zhou in
<bibRefCitation id="EF874B4FAF55FFC1FD9553709C20035E" author="Zhou, X. - M. &amp; Zhang, L. - B." box="[537,827,1894,1918]" pageId="0" pageNumber="155" pagination="630 - 684" refId="ref3031" refString="Zhou, X. - M. &amp; Zhang, L. - B. (2023) Phylogeny, character evolution, and classification of Selaginellaceae. Plant Diversity 45: 630 - 684. https: // doi. org / 10.1016 / j. pld. 2023.07.003" type="journal article" year="2023">Zhou &amp; Zhang (2023: 664)</bibRefCitation>
by
<bibRefCitation id="EF874B4FAF55FFC1FCEE53709CE7035E" author="Alston, A. H. G." box="[866,1020,1894,1918]" pageId="0" pageNumber="155" pagination="41 - 62" refId="ref1600" refString="Alston, A. H. G. (1934 a) The genus Selaginella in the Malay Peninsula (with index). The Gardens Bulletin, Straits Settlements 8: 41 - 62." type="journal article" year="1934">Alston (1934a</bibRefCitation>
, b) and accepted by
<bibRefCitation id="EF874B4FAF55FFC1FB5B53709ABC035E" author="Zhang, X. - C. &amp; Nooteboom, H. P. &amp; Kato, M." box="[1239,1447,1894,1918]" pageId="0" pageNumber="155" pagination="37 - 66" refId="ref2677" refString="Zhang, X. - C., Nooteboom, H. P. &amp; Kato, M. (2013) Selaginellaceae. In: Wu, Z. - Y., Raven, P. H. &amp; Hong, D. - Y. (Eds.) Flora of China, vols. 2 - 3. Science Press, Beijing; Missouri Botanical Garden Press, St. Louis, pp. 37 - 66." type="book chapter" year="2013">
Zhang
<emphasis id="B962EAACAF55FFC1FAAD53719A4C035E" box="[1313,1367,1894,1918]" italics="true" pageId="0" pageNumber="155">et al.</emphasis>
(2013)
</bibRefCitation>
,
<bibRefCitation id="EF874B4FAF55FFC1FF04539C9EB50382" author="Fraser-Jenkins, C. R. &amp; Gandhi, K. N. &amp; Kholia, B. S. &amp; Benniamin, A." box="[136,430,1930,1954]" pageId="0" pageNumber="155" refId="ref1808" refString="Fraser-Jenkins, C. R., Gandhi, K. N., Kholia, B. S. &amp; Benniamin, A. (2017) An annotated checklist of Indian Pteridophytes, Part I (Lycopodiaceae to Thelypteridace). Bishen Singh Mahendra Pal Singh, Dehra Dun, 562 pp." type="book" year="2017">
Fraser-Jenkins
<emphasis id="B962EAACAF55FFC1FEA5539D9E450382" box="[297,350,1930,1954]" italics="true" pageId="0" pageNumber="155">et al.</emphasis>
(2017)
</bibRefCitation>
.
<bibRefCitation id="EF874B4FAF55FFC1FE34539C9DBE0382" author="Shalimov, A. P. &amp; Zaikov, V. F. &amp; Kalyuzhny, S. S. &amp; Zhang, M. - H. &amp; Zhang, X. - C." box="[440,677,1930,1954]" pageId="0" pageNumber="155" pagination="45 - 54" refId="ref2075" refString="Shalimov, A. P., Zaikov, V. F., Kalyuzhny, S. S., Zhang, M. - H. &amp; Zhang, X. - C. (2024) The recovery of Selaginella flaccida based on morphological and molecular data, with the description of S. khokhrjakovii from Indo-China and Peninsular Malaysia. Botanica Pacifica 13: 45 - 54. https: // doi. org / 10.17581 / bp. 2024.13105" type="journal article" year="2024">
Shalimov
<emphasis id="B962EAACAF55FFC1FDA8539D9D420382" box="[548,601,1930,1954]" italics="true" pageId="0" pageNumber="155">et al.</emphasis>
(2024)
</bibRefCitation>
found that the two species differ from each other in leaf morphology and micromorphological features of mega- and microspores and their distinction was corroborated by molecular evidence. We here accept their treatment and recognize
<taxonomicName id="4C164D3DAF55FFC1FDCA53C59DA703CA" authorityName="Li Bing Zhang &amp; X. M. Zhou" authorityYear="2024" baseAuthorityName="Li Bing Zhang &amp; X. M. Zhou" baseAuthorityYear="2024" box="[582,700,2002,2026]" class="Lycopodiopsida" family="Selaginellaceae" genus="Didiclis" kingdom="Plantae" order="Selaginellales" pageId="0" pageNumber="155" phylum="Tracheophyta" rank="species" species="flaccida">
<emphasis id="B962EAACAF55FFC1FDCA53C59DA703CA" box="[582,700,2002,2026]" italics="true" pageId="0" pageNumber="155">D. flaccida</emphasis>
</taxonomicName>
.
</paragraph>
</subSubSection>
</treatment>
</document>

View file

@ -1,41 +1,43 @@
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@ -82,7 +84,7 @@ The island of Lanzarote can be added to the list of the distribution of
in the Canary archipelago: On
<date id="D269100F5B06FF8CFEABF83EFE32F811" box="[353,496,1949,1974]" pageId="0" pageNumber="199" value="2021-11-28">28-11-2021</date>
, a nymph (
<figureCitation id="3EEC2A4A5B06FF8CFDB5F83EFD08F811" box="[639,714,1949,1974]" captionStart="Fig" captionStartId="1.[143,178,1993,2013]" captionTargetId="figure-313@1.[826,1358,1307,1894]" captionTargetPageId="1" captionText="Fig. 2. Nymph of Acrosternum heegeri Fieber, 1861 (Heteroptera: Pentatomidae), Playa Blanca, Lanzarote, Canary Islands, Spain. (Photo: James Senn)." pageId="0" pageNumber="199">Fig. 2</figureCitation>
<figureCitation id="3EEC2A4A5B06FF8CFDB5F83EFD08F811" box="[639,714,1949,1974]" captionStart="Fig" captionStartId="1.[143,178,1993,2013]" captionTargetId="figure-313@1.[826,1358,1307,1894]" captionTargetPageId="1" captionText="Fig. 2. Nymph of Acrosternum heegeri Fieber, 1861 (Heteroptera: Pentatomidae), Playa Blanca, Lanzarote, Canary Islands, Spain. (Photo: James Senn)." figureDoi="http://doi.org/10.5281/zenodo.14947478" httpUri="https://zenodo.org/record/14947478/files/figure.png" pageId="0" pageNumber="199">Fig. 2</figureCitation>
) was found in Playa Blanca, located in the south of the island. The nymph was found behind an apartment in the Elba Hotel complex (James Senn, pers. comm.). A photograph of the specimen was uploaded to the online database iNaturalist (see here).
</emphasis>
</paragraph>

View file

@ -0,0 +1,189 @@
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specimen 4
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<paragraph id="3993EB2F89549BDE022F289EF3086017">
<heading id="D7AC0332FCEC3588F4347048A52855B5" reason="title">Specimen.</heading>
</paragraph>
<paragraph id="96FC032A73715EF0D77ACD7F9096CDDE">
Geoscientific Collection of the University of Göttingen,
<collectingCountry id="A4FFAB4D13BBD06B6CB7F2B412B052CD" name="Germany">Germany</collectingCountry>
(
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),
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.BST.22085
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; syninclusion
<taxonomicName id="93EDEC64065267A4C8D5D7955C81FB01" authorityName="Mamontov, K. Feldberg, Schaf. - Verw., Gradst." authorityYear="2025" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="chiapasensis">
<emphasis id="36AB99B32AB946C050B6A34DAFD76412" italics="true">Frullania chiapasensis</emphasis>
</taxonomicName>
.
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<paragraph id="FE727A74FB749E025F8AEE73F4757DC0">
<heading id="D4E52EB050EEA730FE57DF7684B75EF4" reason="title">Age and stratigraphic level.</heading>
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<paragraph id="FA49282A0298A23F32719E45EBD2AA81">
1523 Ma, Langhian Aquitanian (early to middle Miocene), Simojovel,
<collectingRegion id="63D61821E3F7A89FC0FF37F853F47754" country="Mexico" name="Chiapas">Chiapas</collectingRegion>
,
<collectingCountry id="E03731F95B5BB8221758B10976D4F33E" name="Mexico">Mexico</collectingCountry>
.
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<paragraph id="72AF0339222A8DBE0D6BDD626CA65FCF">
<heading id="1B38AF30626478993617AEC3D264D7A7" reason="title">Description.</heading>
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<paragraph id="9AE33FC12D90BE5047931E6E5F1A7090">
Lichen fragment approximately 2.3 ×
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<emphasis id="9E8F624C4B13BDC3ACDB15F204FB725F" bold="true" italics="true">Thallus</emphasis>
foliose, lobate.
<emphasis id="2BDF7B61F0737E1DC5739F93055F124F" bold="true" italics="true">Lobes</emphasis>
flat and linear, 0.5-1.0 mm wide, lobe apices truncate to slightly rounded (Fig.
<figureCitation id="1B868EDB0046E839EA16C8A9DCDEBD4A" captionStart="Figure 8" captionStartId="F8" captionText="Figure 8. Parmotrema fossils from Dominican and Mexican ambers. A-C. Parmotrema specimen 3 (AMNH DR- 14-294) from Dominican amber; A. Maculate upper surface and long, branching marginal cilia; B. Lower surface with rhizines; C. Dark lower cortex and a cross-section of the lobe revealing the medulla; D, E. Parmotrema specimen 4 (GZG. BST. 22085) from Mexican amber; D. Smooth upper surface and long, dark marginal cilia. The dark, robust rhizines are also visible from the upper side through the semitransparent thallus; E. Branching rhizines on the lower surface." figureDoi="10.3897/fr.28.137758.figure8" httpUri="https://binary.pensoft.net/fig/1264820">8 D</figureCitation>
).
<emphasis id="543BAE8788498B24235F6E83F245A3BD" bold="true" italics="true">Upper surface</emphasis>
smooth, brown (Fig.
<figureCitation id="5FC6ACF41C28E6FFA0D21D032CDC9C8C" captionStart="Figure 8" captionStartId="F8" captionText="Figure 8. Parmotrema fossils from Dominican and Mexican ambers. A-C. Parmotrema specimen 3 (AMNH DR- 14-294) from Dominican amber; A. Maculate upper surface and long, branching marginal cilia; B. Lower surface with rhizines; C. Dark lower cortex and a cross-section of the lobe revealing the medulla; D, E. Parmotrema specimen 4 (GZG. BST. 22085) from Mexican amber; D. Smooth upper surface and long, dark marginal cilia. The dark, robust rhizines are also visible from the upper side through the semitransparent thallus; E. Branching rhizines on the lower surface." figureDoi="10.3897/fr.28.137758.figure8" httpUri="https://binary.pensoft.net/fig/1264820">8 D</figureCitation>
).
<emphasis id="BEBCAEB67DA80052CBF206E252FADC91" bold="true" italics="true">Marginal cilia</emphasis>
dark, mainly simple, up to
<quantity id="66AAF441FE1868FA1C1DD60688B959D3" metricMagnitude="-4" metricUnit="m" metricValue="5.0" unit="mm" value="0.5">0.5 mm</quantity>
long (Fig.
<figureCitation id="1FB63030B27F9E2125F59A05AB8F072B" captionStart="Figure 8" captionStartId="F8" captionText="Figure 8. Parmotrema fossils from Dominican and Mexican ambers. A-C. Parmotrema specimen 3 (AMNH DR- 14-294) from Dominican amber; A. Maculate upper surface and long, branching marginal cilia; B. Lower surface with rhizines; C. Dark lower cortex and a cross-section of the lobe revealing the medulla; D, E. Parmotrema specimen 4 (GZG. BST. 22085) from Mexican amber; D. Smooth upper surface and long, dark marginal cilia. The dark, robust rhizines are also visible from the upper side through the semitransparent thallus; E. Branching rhizines on the lower surface." figureDoi="10.3897/fr.28.137758.figure8" httpUri="https://binary.pensoft.net/fig/1264820">8 D</figureCitation>
).
<emphasis id="735F8CD8F22C4F4ACACE5129DA1CCCF2" bold="true" italics="true">Medulla</emphasis>
not visible.
<emphasis id="DA07FBF4F08B4C8EEDC3698556AF2704" bold="true" italics="true">Lower cortex</emphasis>
concolorous with the upper surface (Fig.
<figureCitation id="65A1C10126D63A0E87B55C118245B1A9" captionStart="Figure 8" captionStartId="F8" captionText="Figure 8. Parmotrema fossils from Dominican and Mexican ambers. A-C. Parmotrema specimen 3 (AMNH DR- 14-294) from Dominican amber; A. Maculate upper surface and long, branching marginal cilia; B. Lower surface with rhizines; C. Dark lower cortex and a cross-section of the lobe revealing the medulla; D, E. Parmotrema specimen 4 (GZG. BST. 22085) from Mexican amber; D. Smooth upper surface and long, dark marginal cilia. The dark, robust rhizines are also visible from the upper side through the semitransparent thallus; E. Branching rhizines on the lower surface." figureDoi="10.3897/fr.28.137758.figure8" httpUri="https://binary.pensoft.net/fig/1264820">8 E</figureCitation>
).
<emphasis id="A5BA0E58680D723FADEAB85059DD6B3E" bold="true" italics="true">Rhizines</emphasis>
dark, abundantly branched (Fig.
<figureCitation id="E1933709A3D28EE7E9E0ADEBCF120E00" captionStart="Figure 8" captionStartId="F8" captionText="Figure 8. Parmotrema fossils from Dominican and Mexican ambers. A-C. Parmotrema specimen 3 (AMNH DR- 14-294) from Dominican amber; A. Maculate upper surface and long, branching marginal cilia; B. Lower surface with rhizines; C. Dark lower cortex and a cross-section of the lobe revealing the medulla; D, E. Parmotrema specimen 4 (GZG. BST. 22085) from Mexican amber; D. Smooth upper surface and long, dark marginal cilia. The dark, robust rhizines are also visible from the upper side through the semitransparent thallus; E. Branching rhizines on the lower surface." figureDoi="10.3897/fr.28.137758.figure8" httpUri="https://binary.pensoft.net/fig/1264820">8 E</figureCitation>
). Apothecia, isidia, soredia, or pycnidia not present.
</paragraph>
</subSubSection>
<subSubSection id="SECID0EDZBI" type="discussion">
<paragraph id="F8F5F0F908F3510A07D8C2104184A732">
<heading id="6657C0923E49AF7011B5CB73944904B6" reason="title">Discussion.</heading>
</paragraph>
<paragraph id="BAC77B90220B66C9F5DC5A6178E21B44">
The general habit and long, tapering marginal cilia identify the specimen as
<taxonomicName id="96E6ED0BDD4B23DE98CE3C14EBA47DBC" class="Lecanoromycetes" family="Parmeliaceae" genus="Parmotrema" kingdom="Fungi" order="Lecanorales" phylum="Ascomycota" rank="genus">
<emphasis id="7692B70178045467D6C5732E76894455" italics="true">Parmotrema</emphasis>
</taxonomicName>
. The branching rhizines and long, tapered marginal cilia resemble the
<taxonomicName id="5226020CA55B8B8704E8468A71DE534D" baseAuthorityName="Elix" baseAuthorityYear="1993" class="Lecanoromycetes" family="Parmeliaceae" genus="Canomaculina" kingdom="Fungi" order="Lecanorales" phylum="Ascomycota" rank="genus">
<emphasis id="F11430B93511594DE3C6D050F6F3CD35" italics="true">Canomaculina</emphasis>
</taxonomicName>
group. However, unlike the
<taxonomicName id="F88C058BC53E24AE01CD63B965F3D28E" baseAuthorityName="Elix" baseAuthorityYear="1993" class="Lecanoromycetes" family="Parmeliaceae" genus="Canomaculina" kingdom="Fungi" order="Lecanorales" phylum="Ascomycota" rank="genus">
<emphasis id="E59A030FC86B86303B1963D08EF06FB1" italics="true">Canomaculina</emphasis>
</taxonomicName>
group, the fossil inclusion seems to possess a smooth upper surface of the lobes that visibly lack maculae.
</paragraph>
</subSubSection>
</treatment>
</document>

View file

@ -0,0 +1,816 @@
<document id="46436C199AFAC5C215E53B98B45C9D20" ID-DOI="10.3897/fr.28.137758" ID-ZooBank="A10C1262-F859-4004-A83C-C8F032C1156A" ID-publisher-id="137758" URI-arpha="66B09103-4195-59B0-BBF5-6B2263A772A9" URI-zoobank="http://zoobank.org/A10C1262-F859-4004-A83C-C8F032C1156A" XM.bibliography_approvedBy="admin" XM.materialsCitations_approvedBy="admin" XM.taxonomicNames_approvedBy="admin" article-type="research-article" checkinTime="1740755594576" checkinUser="pensoft" docAuthor="Feldberg, Kathrin, Kaasalainen, Ulla, Mamontov, Yuriy S., Gradstein, S. Robbert, Schäfer-Verwimp, Alfons, Divakar, Pradeep K. &amp; Schmidt, Alexander R." docDate="2025" docId="B281A1F3C85D593A9770E2E91C0D6D7B" docLanguage="en" docName="Fossil Record 28(1): 79-102" docOrigin="Fossil Record 28 (1)" docSource="https://fr.pensoft.net/article/137758/download/xml/" docStyle="DocumentStyle:PensoftTaxPub.0000.journal_article.generic" docStyleName="PensoftTaxPub.0000.journal_article.generic" docTitle="Frullania chiapasensis Mamontov, K. Feldberg, Schaf. - Verw., Gradst. 2025, sp. nov." docType="treatment" docVersion="2" dtd-version="3.0" lastPageNumber="102" masterDocId="66B09103419559B0BBF56B2263A772A9" masterDocTitle="Extending the fossil record of Miocene neotropical epiphyte communities" masterLastPageNumber="102" masterPageNumber="79" pageNumber="79" updateTime="1740768570675" updateUser="admin">
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<mods:title id="435FEF2E36A327396B681B7551B1751D">Extending the fossil record of Miocene neotropical epiphyte communities</mods:title>
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<mods:namePart id="DEA5178697F82F4BC0F3DBE220CE67AA">Feldberg, Kathrin</mods:namePart>
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<mods:affiliation id="9ADF1D0D30AB2A6D033EADCE74C99937">Department of Geobiology, University of Göttingen, Goldschmidtstraße 3, 37077 Göttingen, Germany</mods:affiliation>
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<mods:affiliation id="7D9D0098F40F7DA382C9C0B119D89915">Tsitsin Main Botanical Garden, Russian Academy of Sciences, Moscow, 127276 Russia</mods:affiliation>
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<mods:affiliation id="6F731C5ADF4378B31F83D4E37B433C17">Departamento de Biología Vegetal II, Facultad de Farmacia, Universidad Complutense de Madrid, 28040 Madrid, Spain</mods:affiliation>
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<mods:affiliation id="083A881D10FF91B84B2CFC6076C4E59C">Department of Geobiology, University of Göttingen, Goldschmidtstraße 3, 37077 Göttingen, Germany</mods:affiliation>
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<mods:title id="182AF499EE3B0679365805BE053E1848">Fossil Record</mods:title>
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<mods:part id="C422BC2DE9072A8348C3B5DC0E7031FE">
<mods:date id="8C068CCF0CA8EEBCABD942891F96864D">2025</mods:date>
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<mods:identifier id="4CB3119920F476ADE4E774A521C4BD20" type="ZooBank">A10C1262-F859-4004-A83C-C8F032C1156A</mods:identifier>
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<treatment id="B281A1F3C85D593A9770E2E91C0D6D7B" ID-arpha="B281A1F3-C85D-593A-9770-E2E91C0D6D7B" ID-zoobank="https://zoobank.org/452DB510-4C0D-4CD4-88BF-C7ACF1EAA9F1" LSID="urn:lsid:plazi:treatment:B281A1F3C85D593A9770E2E91C0D6D7B" LSID-ZBK="urn:lsid:zoobank.org:act:452DB510-4C0D-4CD4-88BF-C7ACF1EAA9F1" httpUri="http://treatment.plazi.org/id/B281A1F3C85D593A9770E2E91C0D6D7B" scope_class="Jungermanniopsida" scope_family="Frullaniaceae" scope_order="Porellales" scope_phylum="Marchantiophyta" scope_subClass="Jungermanniidae">
<subSubSection id="E83500603C50E4CAE9A8FA226A109485" type="nomenclature">
<paragraph id="9FCE6AA9312E18CB8E08D5B02269F5A9">
<taxonomicName id="D5FF49838ADFE6560D6E5BAFB852EAD2" authority="Mamontov, K. Feldberg, Schaf.-Verw., Gradst." authorityName="Mamontov, K. Feldberg, Schaf. - Verw., Gradst." authorityYear="2025" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="chiapasensis" status="sp. nov.">
<emphasis id="5E6BED220A43B10C19534673E01247A7" italics="true">Frullania chiapasensis</emphasis>
Mamontov, K. Feldberg, Schäf.-Verw., Gradst.
</taxonomicName>
<taxonomicNameLabel id="E20D1DDA9EA736A4129A6C935259B12F" rank="species">sp. nov.</taxonomicNameLabel>
</paragraph>
</subSubSection>
<subSubSection id="SECID0EQPAE" type="Holotype">
<paragraph id="0606CB584818E63E4E7BC58542FCDC46">
<heading id="1C9106397B06A99E5ECD24D0615C1CC8" reason="title">
<typeStatus id="F1BE786BBC4502D86FB2EBD3D7ACC3DA" type="holotype">Holotype</typeStatus>
.
</heading>
</paragraph>
<paragraph id="2A6CE8D01260B197729767D51262BE74">
<materialsCitation id="B29BAF6C3078A34FCCDA4096F5D9B1AD" collectionCode="GZG" specimenCode="GZG.BST.22085" specimenCount="1" typeStatus="holotype">
Geoscientific Collection of the University of Göttingen, Germany (
<abbrev content-type="institution" id="ABBRID0EWPAE" xlink_title="Geoscientific Collection of the University of Göttingen">
<collectionCode id="4F13893BEC28ADC6716EC58F239A26B5">GZG</collectionCode>
</abbrev>
),
<specimenCode id="992FA9ED0FA16FBC294FEF0B476DA291">
<abbrev content-type="institution" id="ABBRID0E2PAE" xlink_title="Geoscientific Collection of the University of Göttingen">GZG</abbrev>
.BST.22085
</specimenCode>
; syninclusion
<taxonomicName id="F2DC0EED30F307A11F6599D91F9E7D91" class="Lecanoromycetes" family="Parmeliaceae" genus="Parmotrema" kingdom="Fungi" order="Lecanorales" phylum="Ascomycota" rank="genus">
<emphasis id="897D72B9FE3DDF99B88191E19F6D47BA" italics="true">Parmotrema</emphasis>
</taxonomicName>
specimen 4.
</materialsCitation>
</paragraph>
</subSubSection>
<subSubSection id="SECID0EHQAE" type="etymology">
<paragraph id="7909E72B35A5B68CC4C0089024D8EBA6">
<heading id="6999FFCDF1951C07EEE7FB96BAAFF159" reason="title">Etymology.</heading>
</paragraph>
<paragraph id="8155D1910B4A21644734A736146CF670">
The species is named after the location of the amber deposit in
<collectingRegion id="F1936F1106CE82B193BCD34FA4130A4A" country="Mexico" name="Chiapas">Chiapas</collectingRegion>
,
<collectingCountry id="2DB2EE1E3B4E8354244A44F9F38630F9" name="Mexico">Mexico</collectingCountry>
.
</paragraph>
</subSubSection>
<subSubSection id="SECID0EMQAE" type="Age and stratigraphic level">
<paragraph id="2903A2F8AE7B319A8AAB0D813F41A48B">
<heading id="A141C51B90443A545F6BA3CCE9D445AC" reason="title">Age and stratigraphic level.</heading>
</paragraph>
<paragraph id="0C7CFEB109B4BE86FA9D5E7AEF60F646">
1523 Ma, Langhian Aquitanian (early to middle Miocene), Simojovel,
<collectingRegion id="97F24921A58F7CC48946A44836AE2C2C" country="Mexico" name="Chiapas">Chiapas</collectingRegion>
,
<collectingCountry id="6B2D53A2D036C1970865CC428E297194" name="Mexico">Mexico</collectingCountry>
.
</paragraph>
</subSubSection>
<subSubSection id="SECID0E2QAE" type="diagnosis">
<paragraph id="E236C3589FAFED731C83B703A78BC308">
<heading id="E9E1AB5552D5074EE25C7AC06BD81DF9" reason="title">Diagnosis.</heading>
</paragraph>
<paragraph id="B450E2374DFA511E7F8E26C22160EAFC">
Incubously foliated liverwort with conduplicate-trilobed, entire-margined leaves; dorsal lobe obliquely ovate to elliptical, apex rounded to obtuse, lacking ocelli; ventral lobule
<taxonomicName id="CD78FD2C74FB4BB4B30285462E0BACC0" authorityName="Raddi" authorityYear="1818" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="genus">
<emphasis id="36C3DA74759B9037EE89B6E072014BD3" italics="true">Frullania</emphasis>
</taxonomicName>
- type, saccate, adnate to lobe at a distance of ca. 0.8 of the stem width, parallel or somewhat converging to stem and at places leaning against it, clavate to obovate, not constricted above the postical opening, surface smooth, opening extending along the abaxial side of the lobe for ca. 0.320.50 of the lobule length, sickle-shaped; underleaves triangular to obovate, longer than wide, widest in the upper half, bilobed 0.20.3 × their length, upper half of underleaves armed with 2 short teeth on both sides.
</paragraph>
</subSubSection>
<subSubSection id="SECID0EIRAE" type="description">
<paragraph id="DBFC65249E87E60D7A6E912466A7E49B">
<heading id="BE4BEBF1804C1D2B88F5AE9C6798A8EA" reason="title">Description.</heading>
</paragraph>
<paragraph id="D1EC4891D7353D22A6E640DE62B76C52">
Two gametophyte fragments. Intact shoot ca.
<quantity id="CD40D0BFCEFD065C3EBB74C6DB44C69D" metricMagnitude="-3" metricUnit="m" metricValue="2.64" unit="mm" value="2.64">2.64 mm</quantity>
long,
<quantity id="B5AD407321CF0D17368E988C2329E02E" metricMagnitude="-4" metricUnit="m" metricValue="3.3999999999999995" metricValueMax="3.5" metricValueMin="3.3" unit="mm" value="0.33999999999999997" valueMax="0.35" valueMin="0.33">0.330.35 mm</quantity>
wide with leaves, dark reddish brown (Figs
<figureCitation id="8FDE17F3158DE65AE896D9F843E8844C" captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Frullania chiapasensis sp. nov. from Mexican amber, holotype (GZG. BST. 22085). A. Shoot fragments in dorsal view; B. Shoot fragments in ventral view; C. Detail of intact shoot fragment in ventral view; D. Dorsal lobe (right) with mammillose cells and ventral lobules (left) in dorsal view; E. A toothed underleaf and a lobule with a triangular stylus (indicated by arrow) in ventral view; F. Lobe cells with large central mammilla." figureDoi="10.3897/fr.28.137758.figure1" httpUri="https://binary.pensoft.net/fig/1264813">1 A, B</figureCitation>
,
<figureCitation id="2A6CCCF6D1405DCF1B8D7E573AD44792" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Frullania chiapasensis sp. nov. from Mexican amber, holotype (GZG. BST. 22085). A. Upper part of intact shoot fragment in ventral view; B. Lower part of intact shoot fragment in ventral view, the uppermost underleaf shows a rhizoid bundle." figureDoi="10.3897/fr.28.137758.figure2" httpUri="https://binary.pensoft.net/fig/1264814">2</figureCitation>
); with three broken, lateral branches [not clearly visible].
<emphasis id="0C982543BA39BCA5AABF8BB83E8A0959" bold="true" italics="true">Stem</emphasis>
only slightly darker in color than leaves, 3545 µm in diameter, surface cells not visible.
<emphasis id="3766474B21BF3A66AD451CEA6185BFB2" bold="true" italics="true">Lateral leaves</emphasis>
incubous, alternate, contiguous to imbricate near apex, widely spreading, conduplicate-trilobed (Figs
<figureCitation id="367A7CD146BA74F736344976B263A845" captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Frullania chiapasensis sp. nov. from Mexican amber, holotype (GZG. BST. 22085). A. Shoot fragments in dorsal view; B. Shoot fragments in ventral view; C. Detail of intact shoot fragment in ventral view; D. Dorsal lobe (right) with mammillose cells and ventral lobules (left) in dorsal view; E. A toothed underleaf and a lobule with a triangular stylus (indicated by arrow) in ventral view; F. Lobe cells with large central mammilla." figureDoi="10.3897/fr.28.137758.figure1" httpUri="https://binary.pensoft.net/fig/1264813">1 A C</figureCitation>
,
<figureCitation id="B791B6CE3BB1EAB0407DC3E65197E2C0" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Frullania chiapasensis sp. nov. from Mexican amber, holotype (GZG. BST. 22085). A. Upper part of intact shoot fragment in ventral view; B. Lower part of intact shoot fragment in ventral view, the uppermost underleaf shows a rhizoid bundle." figureDoi="10.3897/fr.28.137758.figure2" httpUri="https://binary.pensoft.net/fig/1264814">2 A</figureCitation>
).
<emphasis id="B4811E963C059D13347DAC67DDDE85EE" bold="true" italics="true">Dorsal lobe</emphasis>
in dorsal aspect convex to nearly flat, partially with reflexed margin, obliquely ovate to elliptical (Figs
<figureCitation id="374F0089B342BAAE298F0C6440A120CC" captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Frullania chiapasensis sp. nov. from Mexican amber, holotype (GZG. BST. 22085). A. Shoot fragments in dorsal view; B. Shoot fragments in ventral view; C. Detail of intact shoot fragment in ventral view; D. Dorsal lobe (right) with mammillose cells and ventral lobules (left) in dorsal view; E. A toothed underleaf and a lobule with a triangular stylus (indicated by arrow) in ventral view; F. Lobe cells with large central mammilla." figureDoi="10.3897/fr.28.137758.figure1" httpUri="https://binary.pensoft.net/fig/1264813">1 A D</figureCitation>
,
<figureCitation id="3FC03B97F189FF1A069F5F0D26621FC8" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Frullania chiapasensis sp. nov. from Mexican amber, holotype (GZG. BST. 22085). A. Upper part of intact shoot fragment in ventral view; B. Lower part of intact shoot fragment in ventral view, the uppermost underleaf shows a rhizoid bundle." figureDoi="10.3897/fr.28.137758.figure2" httpUri="https://binary.pensoft.net/fig/1264814">2 A</figureCitation>
), 150200 µm long × 120150 µm wide, slightly longer than wide, length: width ratio ca. 1.21.3: 1; margin entire, apex rounded to obtuse, inner margin barely extending beyond the farther edge of the stem; lobe base not clearly visible.
<emphasis id="AC38E2B60B3CC4B46E915DACF79A9739" bold="true" italics="true">Lobe cells</emphasis>
almost isodiametric or rectangular to 56 - angled (Fig.
<figureCitation id="E9DC99B968313E161D4C4EC71EB193C2" captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Frullania chiapasensis sp. nov. from Mexican amber, holotype (GZG. BST. 22085). A. Shoot fragments in dorsal view; B. Shoot fragments in ventral view; C. Detail of intact shoot fragment in ventral view; D. Dorsal lobe (right) with mammillose cells and ventral lobules (left) in dorsal view; E. A toothed underleaf and a lobule with a triangular stylus (indicated by arrow) in ventral view; F. Lobe cells with large central mammilla." figureDoi="10.3897/fr.28.137758.figure1" httpUri="https://binary.pensoft.net/fig/1264813">1 D, F</figureCitation>
), 1520 × 1020 µm in the middle of the lobe, marginal cells somewhat smaller; cell walls slightly thickened, with indistinct trigones, no intermediate thickenings seen; with one central mammilla, ca. 5 µm in diameter.
<emphasis id="EDE583B1F60068B36248DC9B5ED3B0B2" bold="true" italics="true">Ocelli</emphasis>
not observed.
<emphasis id="D5E80B30E10A73EDEB1B4393E2F19050" italics="true">
<emphasis id="D33B66E0B964C1D47402999DC4D3A97E" bold="true" italics="true">Ventral lobules</emphasis>
<taxonomicName id="DC40B9A3C988DCA3D1F74FA6186503C9" authorityName="Raddi" authorityYear="1818" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="genus">Frullania</taxonomicName>
</emphasis>
- type, inflated, saccate, explanate lobules not observed; mostly + / - parallel with the stem, isolated lobules obliquely spreading at an angle of ca. 35 ° to the stem, or leaning against it; remotely inserted, distance between anterior base of lobule and edge of stem ca. 0.8 × the stem width; lobules clavate or obovate (Figs
<figureCitation id="FF40B2D8854D8E5DD47D4301CDC94D6C" captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Frullania chiapasensis sp. nov. from Mexican amber, holotype (GZG. BST. 22085). A. Shoot fragments in dorsal view; B. Shoot fragments in ventral view; C. Detail of intact shoot fragment in ventral view; D. Dorsal lobe (right) with mammillose cells and ventral lobules (left) in dorsal view; E. A toothed underleaf and a lobule with a triangular stylus (indicated by arrow) in ventral view; F. Lobe cells with large central mammilla." figureDoi="10.3897/fr.28.137758.figure1" httpUri="https://binary.pensoft.net/fig/1264813">1 C E</figureCitation>
,
<figureCitation id="34D47025711CFF4CE17F00124B1ECF88" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Frullania chiapasensis sp. nov. from Mexican amber, holotype (GZG. BST. 22085). A. Upper part of intact shoot fragment in ventral view; B. Lower part of intact shoot fragment in ventral view, the uppermost underleaf shows a rhizoid bundle." figureDoi="10.3897/fr.28.137758.figure2" httpUri="https://binary.pensoft.net/fig/1264814">2</figureCitation>
), 110140 µm long × 7590 µm wide at the widest part, longer than wide, length: width ratio 1.51.6: 1; apex broadly rounded or angulate; gradually applanate towards the postical opening, not constricted above it, the portion adjoining the opening dorsiventrally compressed in comparison to the gibbous upper half, pinched together; opening very wide, extending along the abaxial side of the lobe for ca. 0.320.50 × the lobule length, sickle-shaped (Fig.
<figureCitation id="DA8654AD99B68A895EF6B196F9F0FFB6" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Frullania chiapasensis sp. nov. from Mexican amber, holotype (GZG. BST. 22085). A. Upper part of intact shoot fragment in ventral view; B. Lower part of intact shoot fragment in ventral view, the uppermost underleaf shows a rhizoid bundle." figureDoi="10.3897/fr.28.137758.figure2" httpUri="https://binary.pensoft.net/fig/1264814">2</figureCitation>
), the apex of the opening with a discolored, gibbous, protuberant cell; the lobules almost symmetric, or somewhat asymmetric due to the sickle-shaped opening, widest in upper third, rarely near the middle; the valves (free margins of the opening) entire, parallel, equal in size; the lobule surface smooth, the margins of the opening crenulate-sinuate, especially near the apex (i. e., the upper end of the sinus), due to protuberant angles between the pairs of cells and strongly concave external walls of these cells.
<emphasis id="7F3FC95DD6D1B22571A31103F3CB19C8" bold="true" italics="true">Styli</emphasis>
indistinctly visible, triangular (Figs
<figureCitation id="E235314092970BBC00BD87C5EE57F6A5" captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Frullania chiapasensis sp. nov. from Mexican amber, holotype (GZG. BST. 22085). A. Shoot fragments in dorsal view; B. Shoot fragments in ventral view; C. Detail of intact shoot fragment in ventral view; D. Dorsal lobe (right) with mammillose cells and ventral lobules (left) in dorsal view; E. A toothed underleaf and a lobule with a triangular stylus (indicated by arrow) in ventral view; F. Lobe cells with large central mammilla." figureDoi="10.3897/fr.28.137758.figure1" httpUri="https://binary.pensoft.net/fig/1264813">1 D, E</figureCitation>
,
<figureCitation id="68A807BCF01CD2C230476B1D91A64E73" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Frullania chiapasensis sp. nov. from Mexican amber, holotype (GZG. BST. 22085). A. Upper part of intact shoot fragment in ventral view; B. Lower part of intact shoot fragment in ventral view, the uppermost underleaf shows a rhizoid bundle." figureDoi="10.3897/fr.28.137758.figure2" httpUri="https://binary.pensoft.net/fig/1264814">2</figureCitation>
), with expanded base from several cells and at least 6 cells high, with a short apex of 2 cells.
<emphasis id="1F93F98E490B07F35F13FDF0CFA692A9" bold="true" italics="true">Underleaves</emphasis>
1 per leaf pair, distant, transversely inserted, ca. 2.12.5 × the stem width, mostly longer than wide (Figs
<figureCitation id="449F13A3C2795B7B4A3F1828625AFEE4" captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Frullania chiapasensis sp. nov. from Mexican amber, holotype (GZG. BST. 22085). A. Shoot fragments in dorsal view; B. Shoot fragments in ventral view; C. Detail of intact shoot fragment in ventral view; D. Dorsal lobe (right) with mammillose cells and ventral lobules (left) in dorsal view; E. A toothed underleaf and a lobule with a triangular stylus (indicated by arrow) in ventral view; F. Lobe cells with large central mammilla." figureDoi="10.3897/fr.28.137758.figure1" httpUri="https://binary.pensoft.net/fig/1264813">1 C, E</figureCitation>
,
<figureCitation id="8FD3C50263586EF306CA22DD33853F24" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Frullania chiapasensis sp. nov. from Mexican amber, holotype (GZG. BST. 22085). A. Upper part of intact shoot fragment in ventral view; B. Lower part of intact shoot fragment in ventral view, the uppermost underleaf shows a rhizoid bundle." figureDoi="10.3897/fr.28.137758.figure2" httpUri="https://binary.pensoft.net/fig/1264814">2</figureCitation>
), 110140 µm long × 100120 µm wide, triangular to obovate, flat, gradually broadening from a cuneate base; lower two thirds entire-margined, upper third armed on both sides with 12 short teeth of 13 cells; bilobed 0.20.3 × their length, sinus acute-angled, the sinus base broadly rounded; lobes widely triangular, obliquely directed to the stem axis, wider than long, with acute to obtuse apex, inner margins of lobes entire, slightly arcuate.
<emphasis id="7311B64EEE30C49207BF6CDCB7CBEE59" bold="true" italics="true">Rhizoids</emphasis>
possibly in fascicles arising from the upper part of the underleaves (Fig.
<figureCitation id="AEBA1F8778D72A17565BF91D9B331A7A" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Frullania chiapasensis sp. nov. from Mexican amber, holotype (GZG. BST. 22085). A. Upper part of intact shoot fragment in ventral view; B. Lower part of intact shoot fragment in ventral view, the uppermost underleaf shows a rhizoid bundle." figureDoi="10.3897/fr.28.137758.figure2" httpUri="https://binary.pensoft.net/fig/1264814">2 B</figureCitation>
). Asexual reproduction not observed.
<emphasis id="86886196E560D86447352DDD9EF0685D" bold="true">
<emphasis id="AF062F2887BBE5C82CC159D9C4E8E2F8" bold="true" italics="true">Sterile</emphasis>
.
</emphasis>
</paragraph>
<caption id="D9135710A22395BEABB48902EC392445" ID-DOI="10.3897/fr.28.137758.figure1" ID-arpha="A05FE2E9-20A0-551F-A701-071869FCEC08" httpUri="https://binary.pensoft.net/fig/1264813" startId="F1">
<paragraph id="81FC27576AC3CB1FE19016DE243D6132">
<label id="7B3716B3030CA2197A308BBE405D5F99">Figure 1.</label>
</paragraph>
<paragraph id="632767DC583D50D7493563BF39427697">
<taxonomicName id="37173C71D6D39BD4B8378B2B495AE34D" authorityName="Mamontov, K. Feldberg, Schaf. - Verw., Gradst." authorityYear="2025" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="chiapasensis" status="sp. nov.">
<emphasis id="4BB866BF2812E3D85AAEBA249DCD302C" italics="true">Frullania chiapasensis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="5D8CD9DC00B0D8DDF2B7852956ECB03C" rank="species">sp. nov.</taxonomicNameLabel>
from Mexican amber, holotype (
<specimenCode id="1BFBC4FF0B5052B961995B84DC415DA3">
<abbrev content-type="institution" id="ABBRID0E4XAE" xlink_title="Geoscientific Collection of the University of Göttingen">GZG</abbrev>
.BST.22085
</specimenCode>
).
<emphasis id="3E033E1D82513F2D719FACC467EEBBF0" bold="true">A.</emphasis>
Shoot fragments in dorsal view;
<emphasis id="B6B01A89B1223B87C4BBC0D408AD96E1" bold="true">B.</emphasis>
Shoot fragments in ventral view;
<emphasis id="2DF4CF5688472D5CEFA4E20A780FC372" bold="true">C.</emphasis>
Detail of intact shoot fragment in ventral view;
<emphasis id="60285EE2C0535583661E9915691385CE" bold="true">D.</emphasis>
Dorsal lobe (right) with mammillose cells and ventral lobules (left) in dorsal view;
<emphasis id="A47249033FDC2C86AA7B12297EA0C458" bold="true">E.</emphasis>
A toothed underleaf and a lobule with a triangular stylus (indicated by arrow) in ventral view;
<emphasis id="C71833C5F7A18E54C6515844E8B222F3" bold="true">F.</emphasis>
Lobe cells with large central mammilla.
</paragraph>
</caption>
<caption id="A2DE05643EE5C181C541CAC29FAC0A9C" ID-DOI="10.3897/fr.28.137758.figure2" ID-arpha="C62FFC19-A8B8-5456-86B5-DF1EDC8C274E" httpUri="https://binary.pensoft.net/fig/1264814" startId="F2">
<paragraph id="4DEE77C7ED15AA6623467D8D3B4C5CBF">
<label id="5B42ECF6FAA8283FA836E69F28ECD1B0">Figure 2.</label>
</paragraph>
<paragraph id="382BE9EEF67C4C8D3228CDE2B6F62FC0">
<taxonomicName id="5EACABF8665C1B7C9350F8D7B64BEF7C" authorityName="Mamontov, K. Feldberg, Schaf. - Verw., Gradst." authorityYear="2025" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="chiapasensis" status="sp. nov.">
<emphasis id="1BDB607011582B8A909869DC1A50242F" italics="true">Frullania chiapasensis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="3E571C4FA343B88AAE458B2FE0D6FD74" rank="species">sp. nov.</taxonomicNameLabel>
from Mexican amber, holotype (
<specimenCode id="F5722AA2F3B7DB45CF1671538B06CCA3">
<abbrev content-type="institution" id="ABBRID0ED1AE" xlink_title="Geoscientific Collection of the University of Göttingen">GZG</abbrev>
.BST.22085
</specimenCode>
).
<emphasis id="B95D0FF137132E246CF8C41A882D453A" bold="true">A.</emphasis>
Upper part of intact shoot fragment in ventral view;
<emphasis id="0DCA37364D5E76B55D9E6D460708DEF0" bold="true">B.</emphasis>
Lower part of intact shoot fragment in ventral view, the uppermost underleaf shows a rhizoid bundle.
</paragraph>
</caption>
</subSubSection>
<subSubSection id="SECID0ER1AE" type="discussion">
<paragraph id="51B11DA637E00BFA2605F93F45C079BA">
<heading id="309D9632CF6EE6436B8A6E7F892D0938" reason="title">Discussion.</heading>
</paragraph>
<paragraph id="89DD496FD1119C9AB3CAF0EAA2E931F4">
The entire conduplicate-trilobed leaves with an entire-margined dorsal lobe and
<taxonomicName id="5CB2A754CFD738191FA8660FF4D1763D" authorityName="Raddi" authorityYear="1818" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="genus">
<emphasis id="C98570FA35822124369E074F368A5846" italics="true">Frullania</emphasis>
</taxonomicName>
- type lobules (Figs
<figureCitation id="9A5055ACB9B32716180ADC9A6A2185B5" captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Frullania chiapasensis sp. nov. from Mexican amber, holotype (GZG. BST. 22085). A. Shoot fragments in dorsal view; B. Shoot fragments in ventral view; C. Detail of intact shoot fragment in ventral view; D. Dorsal lobe (right) with mammillose cells and ventral lobules (left) in dorsal view; E. A toothed underleaf and a lobule with a triangular stylus (indicated by arrow) in ventral view; F. Lobe cells with large central mammilla." figureDoi="10.3897/fr.28.137758.figure1" httpUri="https://binary.pensoft.net/fig/1264813">1 A C</figureCitation>
,
<figureCitation id="1118D4FE2B59A9E4353FD0C486624D02" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Frullania chiapasensis sp. nov. from Mexican amber, holotype (GZG. BST. 22085). A. Upper part of intact shoot fragment in ventral view; B. Lower part of intact shoot fragment in ventral view, the uppermost underleaf shows a rhizoid bundle." figureDoi="10.3897/fr.28.137758.figure2" httpUri="https://binary.pensoft.net/fig/1264814">2 A</figureCitation>
) as well as the bilobed underleaves (Figs
<figureCitation id="431D3CB36A060A898A811B135356FB5F" captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Frullania chiapasensis sp. nov. from Mexican amber, holotype (GZG. BST. 22085). A. Shoot fragments in dorsal view; B. Shoot fragments in ventral view; C. Detail of intact shoot fragment in ventral view; D. Dorsal lobe (right) with mammillose cells and ventral lobules (left) in dorsal view; E. A toothed underleaf and a lobule with a triangular stylus (indicated by arrow) in ventral view; F. Lobe cells with large central mammilla." figureDoi="10.3897/fr.28.137758.figure1" httpUri="https://binary.pensoft.net/fig/1264813">1 C, E</figureCitation>
,
<figureCitation id="A38F3A90D55428F5E7CECFD4DDC017F4" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Frullania chiapasensis sp. nov. from Mexican amber, holotype (GZG. BST. 22085). A. Upper part of intact shoot fragment in ventral view; B. Lower part of intact shoot fragment in ventral view, the uppermost underleaf shows a rhizoid bundle." figureDoi="10.3897/fr.28.137758.figure2" httpUri="https://binary.pensoft.net/fig/1264814">2</figureCitation>
) allow for a reliable generic assignment.
<taxonomicName id="5C73D830DB363D95D33C8BDE9047B0A5" authorityName="Raddi" authorityYear="1818" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="genus">
<emphasis id="A16B0B600638689292A416BAD2FFAB76" italics="true">Frullania</emphasis>
</taxonomicName>
is a large genus of porellalean leafy liverworts with a center of diversity in tropical and subtropical regions, where it often grows epiphytic in the canopy, but it is also a common element of temperate and arctic zones and can grow under much drier conditions than
<taxonomicName id="091B29F4ECECF182DF4A654364773A97" authorityName="Cavers" authorityYear="1910" class="Jungermanniopsida" family="Lejeuneaceae" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="family">Lejeuneaceae</taxonomicName>
(e. g.,
<bibRefCitation id="3C3DF1D3EB78BB0A9FF2007163ACA317" author="Gradstein" etAl="et al." firstAuthor="Gradstein" journalOrPublisher="Memoirs of the New York Botanical Garden" pagination="1-577" refId="B37" refString="Gradstein SR, Churchill SP, Salazar-Allen N (2001) Guide to the Bryophytes of Tropical America. Memoirs of the New York Botanical Garden 86: 1577." title="Guide to the Bryophytes of Tropical America." volume="86" year="2001">Gradstein et al. 2001</bibRefCitation>
).
</paragraph>
<paragraph id="2E0A76402D54E238A1D3C248F18849C8">
<taxonomicName id="6C10B457856F7C858C21B9571A9C343C" authorityName="Raddi" authorityYear="1818" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="genus">
<emphasis id="F1FACD64BEEC60BB095E477662842B91" italics="true">Frullania</emphasis>
</taxonomicName>
is the most diverse genus found in amber. Four species occur in Cretaceous Kachin amber (
<bibRefCitation id="D801EAA40F77FD5CA074F2F5A51AD75A" DOI="10.11646/bde.43.1.6" author="Feldberg K &amp; Gradstein SR &amp; Gröhn C &amp; Heinrichs J &amp; von Konrat M &amp; Mamontov YS &amp; Renner MAM &amp; Roth M &amp; Schäfer-Verwimp A &amp; Sukkharak P &amp; Schmidt AR" journalOrPublisher="Bryophyte Diversity and Evolution" pagination="14-71" refId="B23" refString="Feldberg K, Gradstein SR, Gröhn C, Heinrichs J, von Konrat M, Mamontov YS, Renner MAM, Roth M, Schäfer-Verwimp A, Sukkharak P, Schmidt AR (2021 a) Checklist of fossil liverworts suitable for calibrating phylogenetic reconstructions. Bryophyte Diversity and Evolution 43: 1471. https://doi.org/10.11646/bde.43.1.6" title="Checklist of fossil liverworts suitable for calibrating phylogenetic reconstructions." volume="43">Feldberg et al. 2021 a</bibRefCitation>
,
<bibRefCitation id="41A03B8204DF9A9D33F55C17CBFAEF0E" DOI="10.1016/j.cretres.2021.104987" author="Feldberg K &amp; Schäfer-Verwimp A &amp; Renner MAM &amp; von Konrat M &amp; Bechteler J &amp; Müller P &amp; Wang Y-D &amp; Schneider H &amp; Schmidt AR" refId="B24" refString="Feldberg K, Schäfer-Verwimp A, Renner MAM, von Konrat M, Bechteler J, Müller P, Wang Y-D, Schneider H, Schmidt AR (2021 b) Liverworts from Cretaceous amber. Cretaceous Research 128: 104987. https://doi.org/10.1016/j.cretres.2021.104987">2021 b</bibRefCitation>
), and 16 species have been described from Cenozoic European and African ambers (
<bibRefCitation id="8BD128D4C3816625077EF1D0A686C46C" DOI="10.1111/jse.12796" author="Bouju" etAl="et al." firstAuthor="Bouju" journalOrPublisher="Journal of Systematics and Evolution" pagination="932-954" refId="B9" refString="Bouju V, Feldberg K, Kaasalainen U, Schäfer-Verwimp A, Hedenäs L, Buck WR, Wang B, Perrichot V, Schmidt AR (2021) Miocene Ethiopian amber: A new source of fossil cryptogams. Journal of Systematics and Evolution 60: 932954. https://doi.org/10.1111/jse.12796" title="Miocene Ethiopian amber: A new source of fossil cryptogams." volume="60" year="2021">Bouju et al. 2021</bibRefCitation>
;
<bibRefCitation id="0F14E679BBA2D62AA034FE777A1373A2" DOI="10.11646/bde.43.1.6" author="Feldberg K &amp; Gradstein SR &amp; Gröhn C &amp; Heinrichs J &amp; von Konrat M &amp; Mamontov YS &amp; Renner MAM &amp; Roth M &amp; Schäfer-Verwimp A &amp; Sukkharak P &amp; Schmidt AR" journalOrPublisher="Bryophyte Diversity and Evolution" pagination="14-71" refId="B23" refString="Feldberg K, Gradstein SR, Gröhn C, Heinrichs J, von Konrat M, Mamontov YS, Renner MAM, Roth M, Schäfer-Verwimp A, Sukkharak P, Schmidt AR (2021 a) Checklist of fossil liverworts suitable for calibrating phylogenetic reconstructions. Bryophyte Diversity and Evolution 43: 1471. https://doi.org/10.11646/bde.43.1.6" title="Checklist of fossil liverworts suitable for calibrating phylogenetic reconstructions." volume="43">Feldberg et al. 2021 a</bibRefCitation>
). To date, only three
<taxonomicName id="B31BC8DA8F44FE46DCAB8DFB708BCD63" authorityName="Raddi" authorityYear="1818" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="genus">
<emphasis id="B343BCD76B2E0CEB952CA21B329ED547" italics="true">Frullania</emphasis>
</taxonomicName>
fossils are known from New World deposits: a small fragment described at subgenus level from Dominican amber (
<bibRefCitation id="A5459D9619BFA07047AD035D4F0BA5B6" DOI="10.11646/bde.31.1.15" author="Heinrichs and Schmidt" firstAuthor="Heinrichs" journalOrPublisher="Tropical Bryology" pagination="91-94" refId="B54" refString="Heinrichs J, Schmidt AR (2010) An inclusion of Frullania subgen. Diastaloba s. l. (Frullaniaceae, Porellales) in Dominican amber. Tropical Bryology 31: 9194. https://doi.org/10.11646/bde.31.1.15" title="An inclusion of Frullania subgen. Diastaloba s. l. (Frullaniaceae, Porellales) in Dominican amber." volume="31" year="2010">Heinrichs and Schmidt 2010</bibRefCitation>
) and two fossils from Mexican amber, including
<taxonomicName id="8B4AA093294F9FC6373A7772EAA65E4B" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="undetermined">
<emphasis id="06BA39E20403FE29FED38C7A6A60AFCD" italics="true">Frullania</emphasis>
sp.
</taxonomicName>
(
<bibRefCitation id="8BF25BC946AC730647684D2EF3983CEA" DOI="10.22201/igl.05437652e.2023.12.2.353" author="Juarez-Martinez" etAl="et al." firstAuthor="Juarez-Martinez" journalOrPublisher="Paleontología Mexicana" pagination="107-113" refId="B68" refString="Juárez-Martínez C, Córdova-Tabares VM, Estrada-Ruiz E (2023) Primer registro de una inclusión de planta hepática del género Frullania (Frullaniaceae, Porellales) en el ámbar miocénico de Chiapas, México. Paleontología Mexicana 12: 107113. https://doi.org/10.22201/igl.05437652e.2023.12.2.353" title="Primer registro de una inclusión de planta hepática del género Frullania (Frullaniaceae, Porellales) en el ámbar miocénico de Chiapas, México." volume="12" year="2023">Juárez-Martínez et al. 2023</bibRefCitation>
) and the recently described
<taxonomicName id="6354D45A516AB1DEC918E1D6DE25011A" authority="Juarez-Mart. &amp; Estrada-Ruiz" authorityName="Juarez-Mart. &amp; Estrada-Ruiz" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="delgadilloi">
<emphasis id="DA2F6A93639BED7331D5335BE03D2FBE" italics="true">F. delgadilloi</emphasis>
Juárez-Mart. &amp; Estrada-Ruiz
</taxonomicName>
, the latter representing the most completely preserved specimen (
<bibRefCitation id="66DF8700BFE44760C3F0832B7F223B5C" DOI="10.1016/j.jsames.2024.104858" author="Juarez-Martinez" etAl="et al." firstAuthor="Juarez-Martinez" refId="B69" refString="Juárez-Martínez C, Córdova-Tabares VM, Estrada-Ruiz E (2024) A new fossil species of a liverwort of the Frullania genus (Frullaniaceae, Marchantiophyta) from the Miocene amber of Simojovel de Allende, Chiapas, Mexico. Journal of South American Earth Sciences 137: 104858. https://doi.org/10.1016/j.jsames.2024.104858" year="2024">Juárez-Martínez et al. 2024</bibRefCitation>
).
<collectingCountry id="C0A7079B27F3B7A4BD3E1B8BC6FABA2B" name="Dominican Republic">The Dominican</collectingCountry>
fossil differs from
<taxonomicName id="744305A1B320B9BA810CDD9770450DE8" authorityName="Mamontov, K. Feldberg, Schaf. - Verw., Gradst." authorityYear="2025" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="chiapasensis">
<emphasis id="256F488EA41D2BE2707D219B4B66BC53" italics="true">F. chiapasensis</emphasis>
</taxonomicName>
in possessing obliquely spreading lobules, small, acuminate styli, and more deeply bilobed, wider than long underleaves (
<bibRefCitation id="420D63C69ED439B21230A4DF25B9E648" DOI="10.11646/bde.31.1.15" author="Heinrichs and Schmidt" firstAuthor="Heinrichs" journalOrPublisher="Tropical Bryology" pagination="91-94" refId="B54" refString="Heinrichs J, Schmidt AR (2010) An inclusion of Frullania subgen. Diastaloba s. l. (Frullaniaceae, Porellales) in Dominican amber. Tropical Bryology 31: 9194. https://doi.org/10.11646/bde.31.1.15" title="An inclusion of Frullania subgen. Diastaloba s. l. (Frullaniaceae, Porellales) in Dominican amber." volume="31" year="2010">Heinrichs and Schmidt 2010</bibRefCitation>
, figs 15).
<taxonomicName id="50872EB5F306CEECD30EC13EECD6201A" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="undetermined">
<emphasis id="BB16674B42B2F8F2B9CEB443F786A14C" italics="true">Frullania</emphasis>
sp.
</taxonomicName>
from Mexican amber is differentiated by its lobules which are not much longer than wide, constricted above the only slightly arched opening, and much smaller in relation to the lobes (
<bibRefCitation id="7D9BD3B0F492A73645A26548DAF3D2F5" DOI="10.22201/igl.05437652e.2023.12.2.353" author="Juarez-Martinez" etAl="et al." firstAuthor="Juarez-Martinez" journalOrPublisher="Paleontología Mexicana" pagination="107-113" refId="B68" refString="Juárez-Martínez C, Córdova-Tabares VM, Estrada-Ruiz E (2023) Primer registro de una inclusión de planta hepática del género Frullania (Frullaniaceae, Porellales) en el ámbar miocénico de Chiapas, México. Paleontología Mexicana 12: 107113. https://doi.org/10.22201/igl.05437652e.2023.12.2.353" title="Primer registro de una inclusión de planta hepática del género Frullania (Frullaniaceae, Porellales) en el ámbar miocénico de Chiapas, México." volume="12" year="2023">Juárez-Martínez et al. 2023</bibRefCitation>
, fig. 3). Moreover, the leaf lobules in this specimen have a crook-shaped mouth similar to those in the subgenus
<taxonomicName id="2A26A94DC476D63BBDB2D7F794D38E85" authority="Spruce. By" authorityName="Spruce. By" class="Bryopsida" family="Meteoriaceae" genus="Meteoriopsis" kingdom="Plantae" order="Leucodontales" phylum="Bryophyta" rank="genus">
<emphasis id="BD6146523586A361AB2C282D6D0D73C3" italics="true">Meteoriopsis</emphasis>
Spruce. By
</taxonomicName>
contrast, in
<taxonomicName id="9706DAD98AFDF3E5A3EC44D6408D9811" authorityName="Mamontov, K. Feldberg, Schaf. - Verw., Gradst." authorityYear="2025" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="chiapasensis">
<emphasis id="A6203BB440ECB44F23ACD3C17E6FBC3A" italics="true">F. chiapasensis</emphasis>
</taxonomicName>
the lobule mouth is sickle-shaped and resembles those of numerous species in the subgenera
<emphasis id="48402354ED54164875AA94FC5AAE2FCE" italics="true">Diastaloba</emphasis>
s. l. and
<taxonomicName id="EE6B69EDF7EDBACF2060D893C6BBD2EC" rank="subGenus" subGenus="Microfrullania">
<emphasis id="81F4E19B486FCB48C0097F69724BD5DA" italics="true">Microfrullania</emphasis>
</taxonomicName>
(R. M. Schust.) R. M. Schust. (see also below). The underleaves of
<taxonomicName id="FF1E0A97EB02700071B7C51AE3959387" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="undetermined">
<emphasis id="2DC7C81F658B6FA33F25D65739E8C1F0" italics="true">Frullania</emphasis>
sp.
</taxonomicName>
are not well visible and cannot be compared.
<taxonomicName id="B073F001E7CC38023C4B2EE1545BA137" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="delgadilloi">
<emphasis id="07B44D39B0D87588A151D53CC2029C29" italics="true">Frullania delgadilloi</emphasis>
</taxonomicName>
differs from
<taxonomicName id="B5DE7D22362E33DBC5DE0CF14EE6925B" authorityName="Mamontov, K. Feldberg, Schaf. - Verw., Gradst." authorityYear="2025" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="chiapasensis">
<emphasis id="6522B182CF7FB77734879EA6B83A2ABF" italics="true">F. chiapasensis</emphasis>
</taxonomicName>
in lobule and underleaf shape. The lobules of
<taxonomicName id="C27AD1C343DAD969BCFFB368C87E479D" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="delgadilloi">
<emphasis id="F7790BFE625E018641E1574E199BF734" italics="true">F. delgadilloi</emphasis>
</taxonomicName>
are significantly smaller in relation to the lobes and strongly oblique with the apices oriented outwards, resembling
<collectingCountry id="E5757C178F5F22609331C7AB1B9E2C37" name="Dominican Republic">the Dominican</collectingCountry>
<taxonomicName id="2E8DB2F7E51421BCC61C0E2A90F7E64E" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="undetermined">
<emphasis id="DED0872AAE8D2E8A5D7175CEF64D1B33" italics="true">Frullania</emphasis>
sp.
</taxonomicName>
in this aspect. Furthermore, the lobules of
<taxonomicName id="D8DA8743563D0C6315AF274B276C9565" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="delgadilloi">
<emphasis id="5FA990991A937BF6D833527E140C9FCE" italics="true">F. delgadilloi</emphasis>
</taxonomicName>
are inserted further from the stem (ca. 12 times the stem width), while those of
<taxonomicName id="D84F7D29228ECBDA206CEB8A5A30DAED" authorityName="Mamontov, K. Feldberg, Schaf. - Verw., Gradst." authorityYear="2025" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="chiapasensis">
<emphasis id="9EE37A3F84EBF604E270B9B77AE4780E" italics="true">F. chiapasensis</emphasis>
</taxonomicName>
are larger in relation to the lobes, less remote (ca. 0.8 times the stem width), and oriented mostly parallel to the stem. The lobes of
<taxonomicName id="F53E77BFAD701D2FD6616F3307F2286F" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="delgadilloi">
<emphasis id="6166E87F521E7309BDA34FF5CE5E5B8C" italics="true">F. delgadilloi</emphasis>
</taxonomicName>
possess scattered larger cells, interpreted by the authors as ocelli, which seem to be absent from
<taxonomicName id="9C2F370B242D8E1EE7ADB3513F6740BA" authorityName="Mamontov, K. Feldberg, Schaf. - Verw., Gradst." authorityYear="2025" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="chiapasensis">
<emphasis id="98DDFBD74D45CA984C4974BBF22487F3" italics="true">F. chiapasensis</emphasis>
</taxonomicName>
. The underleaves in
<taxonomicName id="8A176CBC1FD3E89989FC2F2AB684D486" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="delgadilloi">
<emphasis id="0400F9543802DABE4D0B6246EC106486" italics="true">F. delgadilloi</emphasis>
</taxonomicName>
are ovate, four or more times wider than the stem (according to our measurements of the shoots imaged in the description of
<taxonomicName id="39819BEFFE5E75642F0850714D9338C6" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="delgadilloi">
<emphasis id="C5BB193437884481A96D107C97D7E816" italics="true">F. delgadilloi</emphasis>
</taxonomicName>
), bilobed to ca. 0.3 of their length, with the lobes narrowly acute and angular along the outer margins (
<bibRefCitation id="65B98F1C991A2D04A898858DA047BEA4" DOI="10.1016/j.jsames.2024.104858" author="Juarez-Martinez" etAl="et al." firstAuthor="Juarez-Martinez" refId="B69" refString="Juárez-Martínez C, Córdova-Tabares VM, Estrada-Ruiz E (2024) A new fossil species of a liverwort of the Frullania genus (Frullaniaceae, Marchantiophyta) from the Miocene amber of Simojovel de Allende, Chiapas, Mexico. Journal of South American Earth Sciences 137: 104858. https://doi.org/10.1016/j.jsames.2024.104858" year="2024">Juárez-Martínez et al. 2024</bibRefCitation>
, fig. 3 A,
<collectionCode id="874DD0C5A186E514E8DABD7F00F15681" country="USA" lsid="urn:lsid:biocol.org:col:15707" name="Field Museum of Natural History, Botany Department" type="Herbarium">F</collectionCode>
). In contrast, the underleaves of
<taxonomicName id="384A7E02A39DB2A4D52192AFC55DCB1E" authorityName="Mamontov, K. Feldberg, Schaf. - Verw., Gradst." authorityYear="2025" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="chiapasensis">
<emphasis id="DBA9C3A1E0CC4F72ACE4C1408A9E265B" italics="true">F. chiapasensis</emphasis>
</taxonomicName>
are triangular to obovate, less than three times wider than the stem, and bilobed to 0.20.3 of their length, with 12 short but distinct teeth at the base of the obtuse to acute lobes. However, the species also resemble each other in several ways:
<taxonomicName id="002C557DEECD945C2A801325C7CB2391" authorityName="Mamontov, K. Feldberg, Schaf. - Verw., Gradst." authorityYear="2025" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="chiapasensis">
<emphasis id="BBB3AF274C931C4BF029110C16DA5AB7" italics="true">F. chiapasensis</emphasis>
</taxonomicName>
is characterized by a sickle-shaped lobule mouth that extends to ca. 0.320.50 of the lobule length and has crenulate margins as well as a protuberant cell at the apex.
<bibRefCitation id="5E7A0EC17C631D77E5838B59EAAD39F2" DOI="10.1016/j.jsames.2024.104858" author="Juárez-Martínez C &amp; Córdova-Tabares VM &amp; Estrada-Ruiz E" etAl="et al." firstAuthor="Juarez-Martinez" refId="B69" refString="Juárez-Martínez C, Córdova-Tabares VM, Estrada-Ruiz E (2024) A new fossil species of a liverwort of the Frullania genus (Frullaniaceae, Marchantiophyta) from the Miocene amber of Simojovel de Allende, Chiapas, Mexico. Journal of South American Earth Sciences 137: 104858. https://doi.org/10.1016/j.jsames.2024.104858" year="2024">Juárez-Martínez et al. (2024)</bibRefCitation>
describe the opening as “ asymmetrical slightly crenulate with incision reaching about ca. 0.25 of lobule length ”; a protuberant cell is not mentioned. Furthermore, both species have large, foliose styli. We consider especially the differences in lobule shape and orientation to be significant enough to describe a second
<taxonomicName id="2B853F87797BA4B13A6249E9A8FB9FBE">
<emphasis id="E214D1C3D9B115D09C29B20603FF215F" italics="true">Frullania</emphasis>
species
</taxonomicName>
for Mexican Chiapas amber, though both species might be related at subgenus or section level.
</paragraph>
<paragraph id="3AD9D22BAEF1C92E06A8B99089DCA5EF">
Other liverwort inclusions similar to
<taxonomicName id="7D37A68776D29239DF396D98A52D8373" authorityName="Mamontov, K. Feldberg, Schaf. - Verw., Gradst." authorityYear="2025" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="chiapasensis">
<emphasis id="713CEAC1AC795D05A7148494598F3C6C" italics="true">F. chiapasensis</emphasis>
</taxonomicName>
are
<taxonomicName id="35BF8C1DE3946FA33D5EB49A39C09149" authority="Grolle" authorityName="Grolle" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="baltica">
<emphasis id="E38C1E2709E2B1915153A56D464ADA57" italics="true">F. baltica</emphasis>
Grolle
</taxonomicName>
from Baltic and Bitterfeld amber and
<taxonomicName id="7E727D454CDED970A38F13CEC7B5347F" authority="Mamontov et al." authorityName="Mamontov" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="schmalhausenii">
<emphasis id="337F10C281119FCE17AAA682D03DAE54" italics="true">F. schmalhausenii</emphasis>
Mamontov et al.
</taxonomicName>
from
<collectingRegion id="79F87EF882B66C4BD28DB97469563D15" country="Ukraine" name="Rivne">Rovno</collectingRegion>
amber. However, these species are not only older than
<taxonomicName id="AC73AF21844F329A2C093E0ED2A7434E" authorityName="Mamontov, K. Feldberg, Schaf. - Verw., Gradst." authorityYear="2025" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="chiapasensis">
<emphasis id="82C2C91A68AD93B78C2A244877B32019" italics="true">F. chiapasensis</emphasis>
</taxonomicName>
but can easily be differentiated based on their lobules, which are obliquely spreading, with the apices oriented away from the stem like in
<taxonomicName id="F8229B45D645DEC55F0261BFCD15FE0C" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="delgadilloi">
<emphasis id="0C5E55D19BB94A128DA98F6E412E6E21" italics="true">F. delgadilloi</emphasis>
</taxonomicName>
, instead of lobules oriented mostly parallel to the stem. They can also be distinguished by their low abaxial lobule opening, which ends at less than 0.1 of the lobule length, while the lobe opening of
<taxonomicName id="CE014D86F27A70B5F638F483F86067FC" authorityName="Mamontov, K. Feldberg, Schaf. - Verw., Gradst." authorityYear="2025" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="chiapasensis">
<emphasis id="26223BA1AA024ECC30E7003DBDF21EB1" italics="true">F. chiapasensis</emphasis>
</taxonomicName>
is strongly extended along the lateral lobe margin and hence sickle-shaped (Figs
<figureCitation id="421F6586494DF3303EE4BA35F4F27EF8" captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Frullania chiapasensis sp. nov. from Mexican amber, holotype (GZG. BST. 22085). A. Shoot fragments in dorsal view; B. Shoot fragments in ventral view; C. Detail of intact shoot fragment in ventral view; D. Dorsal lobe (right) with mammillose cells and ventral lobules (left) in dorsal view; E. A toothed underleaf and a lobule with a triangular stylus (indicated by arrow) in ventral view; F. Lobe cells with large central mammilla." figureDoi="10.3897/fr.28.137758.figure1" httpUri="https://binary.pensoft.net/fig/1264813">1</figureCitation>
,
<figureCitation id="6D9A97E7D2FE1551002143F587C77E28" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Frullania chiapasensis sp. nov. from Mexican amber, holotype (GZG. BST. 22085). A. Upper part of intact shoot fragment in ventral view; B. Lower part of intact shoot fragment in ventral view, the uppermost underleaf shows a rhizoid bundle." figureDoi="10.3897/fr.28.137758.figure2" httpUri="https://binary.pensoft.net/fig/1264814">2</figureCitation>
;
<bibRefCitation id="1411E5F4CFA2477BA7DD4D078493AA42" author="Grolle and Meister" firstAuthor="Grolle" journalOrPublisher="Weissdorn Verlag, Jena" refId="B50" refString="Grolle R, Meister K (2004) The Liverworts in Baltic and Bitterfeld Amber. Weissdorn Verlag, Jena, 91 pp." title="The Liverworts in Baltic and Bitterfeld Amber." year="2004">Grolle and Meister 2004</bibRefCitation>
, plate 5;
<bibRefCitation id="AD81068579AB0DA3EE4192A0470DAD4A" DOI="10.1134/S0031030119100113" author="Mamontov" etAl="et al." firstAuthor="Mamontov" journalOrPublisher="Paleontological Journal" pagination="1095-1103" refId="B87" refString="Mamontov YS, Ignatov MS, Perkovsky EE (2019) Liverworts from Rovno amber (Ukraine). 8. Frullania ekaterinae sp. nov. and F. schmalhausenii sp. nov. Paleontological Journal 53: 10951103. https://doi.org/10.1134/S0031030119100113" title="Liverworts from Rovno amber (Ukraine). 8. Frullania ekaterinae sp. nov. and F. schmalhausenii sp. nov." volume="53" year="2019">Mamontov et al. 2019</bibRefCitation>
, figs 2, 3 b, c, g, j, k).
</paragraph>
<paragraph id="C32914D02DA74BCD095FD3D8BD996FBA">
Because Mexican amber is a rather young deposit and several extant species have been found in Dominican amber (
<bibRefCitation id="E23DA36BFD9AF0DA070245764C499F6A" author="Gradstein" firstAuthor="Gradstein" journalOrPublisher="Nova Hedwigia" pagination="353-374" refId="B31" refString="Gradstein SR (1993) New fossil Hepaticae preserved in amber of the Dominican Republic. Nova Hedwigia 57: 353374." title="New fossil Hepaticae preserved in amber of the Dominican Republic." volume="57" year="1993">Gradstein 1993</bibRefCitation>
;
<bibRefCitation id="183CD31064F02419C2446F7A06927B81" DOI="10.1093/botlinnean/box041" author="Kubilius" etAl="et al." firstAuthor="Kubilius" journalOrPublisher="Botanical Journal of the Linnean Society" pagination="56-64" refId="B76" refString="Kubilius RA, Bölz A, Feldberg K, Hedenäs L, Schäfer-Verwimp A, Schmidt AR, Heinrichs J (2017) The moss Helicophyllum torquatum (Bryopsida: Helicophyllaceae) has survived since at least the Miocene. Botanical Journal of the Linnean Society 185: 5664. https://doi.org/10.1093/botlinnean/box041" title="The moss Helicophyllum torquatum (Bryopsida: Helicophyllaceae) has survived since at least the Miocene." volume="185" year="2017">Kubilius et al. 2017</bibRefCitation>
;
<bibRefCitation id="2E430770714BBDAC5AB2D61CC3510AFA" DOI="10.11646/bde.43.1.6" author="Feldberg K &amp; Gradstein SR &amp; Gröhn C &amp; Heinrichs J &amp; von Konrat M &amp; Mamontov YS &amp; Renner MAM &amp; Roth M &amp; Schäfer-Verwimp A &amp; Sukkharak P &amp; Schmidt AR" journalOrPublisher="Bryophyte Diversity and Evolution" pagination="14-71" refId="B23" refString="Feldberg K, Gradstein SR, Gröhn C, Heinrichs J, von Konrat M, Mamontov YS, Renner MAM, Roth M, Schäfer-Verwimp A, Sukkharak P, Schmidt AR (2021 a) Checklist of fossil liverworts suitable for calibrating phylogenetic reconstructions. Bryophyte Diversity and Evolution 43: 1471. https://doi.org/10.11646/bde.43.1.6" title="Checklist of fossil liverworts suitable for calibrating phylogenetic reconstructions." volume="43">Feldberg et al. 2021 a</bibRefCitation>
), it is very important to compare the new fossil to the extant diversity. As one of the largest and taxonomically most complex genera of leafy liverworts,
<taxonomicName id="0EE65E9C2A39F10F452C768195C43A2C" authorityName="Raddi" authorityYear="1818" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="genus">
<emphasis id="95DB3AEA38ACC7E16B340A4BB5D9A033" italics="true">Frullania</emphasis>
</taxonomicName>
contains more than 2,000 published names (
<bibRefCitation id="C1E58FE749D768FC695F48C037216A84" DOI="10.1127/0029-5035/2010/0091-0471" author="von Konrat" etAl="et al." firstAuthor="von Konrat" journalOrPublisher="Nova Hedwigia" pagination="471-500" refId="B137" refString="von Konrat M, Hentschel J, Heinrichs J, Braggins JE, Pócs T (2010) Forty-one degrees below and sixty years in the dark: Frullania sect. Inconditum, a new section of Australasian Frullania species including F. colliculosa, sp. nov., F. hodgsoniae, nom. and stat. nov., F. aterrima, and F. hattorii (Frullaniaceae, Marchantiophyta). Nova Hedwigia 91: 471500. https://doi.org/10.1127/0029-5035/2010/0091-0471" title="Forty-one degrees below and sixty years in the dark: Frullania sect. Inconditum, a new section of Australasian Frullania species including F. colliculosa, sp. nov., F. hodgsoniae, nom. and stat. nov., F. aterrima, and F. hattorii (Frullaniaceae, Marchantiophyta)." volume="91" year="2010">von Konrat et al. 2010</bibRefCitation>
;
<bibRefCitation id="4F5B92A08F36C297B477C75FF1D35EBA" DOI="10.11646/phytotaxa.220.2.3" author="Hentschel" etAl="et al." firstAuthor="Hentschel" journalOrPublisher="Phytotaxa" pagination="127-142" refId="B63" refString="Hentschel J, von Konrat M, Söderström L, Hagborg A, Larraín J, Sukkharak P, Uribe J, Zhang L (2015) Notes on Early Land Plants Today. 72. Infrageneric classification and new combinations, new names, new synonyms in Frullania (Marchantiophyta). Phytotaxa 220: 127142. https://doi.org/10.11646/phytotaxa.220.2.3" title="Notes on Early Land Plants Today. 72. Infrageneric classification and new combinations, new names, new synonyms in Frullania (Marchantiophyta)." volume="220" year="2015">Hentschel et al. 2015</bibRefCitation>
) and 675 currently accepted taxa (
<bibRefCitation id="1C6CE9B1EE2587ECDBC93DBDA9A02682" DOI="10.3897/phytokeys.59.6261" author="Soderstrom" etAl="et al." firstAuthor="Soderstrom" journalOrPublisher="PhytoKeys" pagination="1-828" refId="B123" refString="Söderström L, Hagborg A, von Konrat M, Bartholomew-Began S, Bell D, Briscoe L, Brown E, Cargill DC, Costa DP, Crandall-Stotler BJ, Cooper ED, Dauphin G, Engel JJ, Feldberg K, Glenny D, Gradstein SR, He X, Heinrichs J, Hentschel J, Ilkiu-Borges AL, Katagiri T, Konstantinova NA, Larraín J, Long DG, Nebel M, Pócs T, Puche F, Reiner-Drehwald E, Renner MAM, Sass-Gyarmati A, Schäfer-Verwimp A, Moragues JGS, Stotler RE, Sukkharak P, Thiers BM, Uribe J, Váňa J, Villarreal JC, Wigginton M, Zhang L, Zhu R-L (2016) World checklist of hornworts and liverworts. PhytoKeys 59: 1828. https://doi.org/10.3897/phytokeys.59.6261" title="World checklist of hornworts and liverworts." volume="59" year="2016">Söderström et al. 2016</bibRefCitation>
;
<bibRefCitation id="6DCDA9A3A6EDA21CEC308D28ABBF24CA" author="Schafer-Verwimp and Winter" firstAuthor="Schafer-Verwimp" journalOrPublisher="Frahmia" pagination="1-14" refId="B114" refString="Schäfer-Verwimp A, Winter G (2023) Frullania hohenesteri (Marchantiophyta: Frullaniaceae) a new species from Santa Catarina, South Brazil. Frahmia 39: 114. http://www.frahmia.de/downloads/frahmia/frahmia_0039.pdf" title="Frullania hohenesteri (Marchantiophyta: Frullaniaceae) - a new species from Santa Catarina, South Brazil." url="http://www.frahmia.de/downloads/frahmia/frahmia_0039.pdf" volume="39" year="2023">Schäfer-Verwimp and Winter 2023</bibRefCitation>
). Many extant
<taxonomicName id="A3D55B7D1007C803A865C99429FB6E2F">
<emphasis id="4363F006A7FD672058C9CF21FCD4FB97" italics="true">Frullania</emphasis>
species
</taxonomicName>
and subspecies are insufficiently known, and information about their morphology is only available in protologues published in the 19 th and early 20 th centuries. Recent studies have greatly expanded the knowledge of the complex taxonomy of the genus, and it seems that its diversity is even greater than anticipated, as new species are found regularly (e. g.,
<bibRefCitation id="FDF1A6A55D12B650ADE6F07D16E82F3C" DOI="10.1016/j.ympev.2008.12.021" author="Hentschel" etAl="et al." firstAuthor="Hentschel" journalOrPublisher="Molecular Phylogenetics and Evolution" pagination="142-156" refId="B62" refString="Hentschel J, von Konrat MJ, Pócs T, Schäfer-Verwimp A, Shaw AJ, Schneider H, Heinrichs J (2009) Molecular insights into the phylogeny and subgeneric classification of Frullania Raddi (Frullaniaceae, Porellales). Molecular Phylogenetics and Evolution 52: 142156. https://doi.org/10.1016/j.ympev.2008.12.021" title="Molecular insights into the phylogeny and subgeneric classification of Frullania Raddi (Frullaniaceae, Porellales)." volume="52" year="2009">Hentschel et al. 2009</bibRefCitation>
,
<bibRefCitation id="6D801407E8513453E0697D21828959D9" DOI="10.11646/phytotaxa.220.2.3" author="Hentschel" firstAuthor="Hentschel" journalOrPublisher="Phytotaxa" pagination="127-142" refId="B63" refString="Hentschel J, von Konrat M, Söderström L, Hagborg A, Larraín J, Sukkharak P, Uribe J, Zhang L (2015) Notes on Early Land Plants Today. 72. Infrageneric classification and new combinations, new names, new synonyms in Frullania (Marchantiophyta). Phytotaxa 220: 127142. https://doi.org/10.11646/phytotaxa.220.2.3" title="Notes on Early Land Plants Today. 72. Infrageneric classification and new combinations, new names, new synonyms in Frullania (Marchantiophyta)." volume="220" year="2015">2015</bibRefCitation>
;
<bibRefCitation id="5D2431B2361881F1F4B5AFFA74894C40" DOI="10.1016/j.ympev.2010.05.004" author="Heinrichs" etAl="et al." firstAuthor="Heinrichs" journalOrPublisher="Molecular Phylogenetics and Evolution" pagination="1105-1114" refId="B56" refString="Heinrichs J, Hentschel J, Bombosch A, Fiebig A, Reise J, Edelmann M, Kreier H-P, Schäfer-Verwimp A, Caspari S, Schmidt AR, Zhu R-L, von Konrat M, Shaw B, Shaw AJ (2010) One species or at least eight? Delimitation and distribution of Frullania tamarisci (L.) Dumort. s. l. (Jungermanniopsida, Porellales) inferred from nuclear and chloroplast DNA markers. Molecular Phylogenetics and Evolution 56: 11051114. https://doi.org/10.1016/j.ympev.2010.05.004" title="One species or at least eight? Delimitation and distribution of Frullania tamarisci (L.) Dumort. s. l. (Jungermanniopsida, Porellales) inferred from nuclear and chloroplast DNA markers." volume="56" year="2010">Heinrichs et al. 2010</bibRefCitation>
;
<bibRefCitation id="67BE891E8BA4A5931DFFB913E09FE0FE" DOI="10.3732/ajb.1000171" author="Ramaiya" etAl="et al." firstAuthor="Ramaiya" journalOrPublisher="American Journal of Botany" pagination="1707-1718" refId="B102" refString="Ramaiya M, Johnson MG, Shaw B, Heinrichs J, Hentschel J, von Konrat M, Davison PG, Shaw AJ (2010) Morphologically cryptic biological species within the liverwort Frullania asagrayana. American Journal of Botany 97: 17071718. https://doi.org/10.3732/ajb.1000171" title="Morphologically cryptic biological species within the liverwort Frullania asagrayana." volume="97" year="2010">Ramaiya et al. 2010</bibRefCitation>
;
<bibRefCitation id="2031EBA89A81E3CD38689B82DFFEFC33" DOI="10.1127/0029-5035/2010/0091-0471" author="von Konrat" etAl="et al." firstAuthor="von Konrat" journalOrPublisher="Nova Hedwigia" pagination="471-500" refId="B137" refString="von Konrat M, Hentschel J, Heinrichs J, Braggins JE, Pócs T (2010) Forty-one degrees below and sixty years in the dark: Frullania sect. Inconditum, a new section of Australasian Frullania species including F. colliculosa, sp. nov., F. hodgsoniae, nom. and stat. nov., F. aterrima, and F. hattorii (Frullaniaceae, Marchantiophyta). Nova Hedwigia 91: 471500. https://doi.org/10.1127/0029-5035/2010/0091-0471" title="Forty-one degrees below and sixty years in the dark: Frullania sect. Inconditum, a new section of Australasian Frullania species including F. colliculosa, sp. nov., F. hodgsoniae, nom. and stat. nov., F. aterrima, and F. hattorii (Frullaniaceae, Marchantiophyta)." volume="91" year="2010">von Konrat et al. 2010</bibRefCitation>
,
<bibRefCitation id="1B716C75C67E22C720FD5998E0022511" DOI="10.3897/phytokeys.8.2496" author="von Konrat" firstAuthor="von Konrat" journalOrPublisher="PhytoKeys" pagination="13-36" refId="B138" refString="von Konrat M, de Lange P, Greif M, Strozier L, Hentschel J, Heinrichs J (2012) Frullania knightbridgei, a new liverwort (Frullaniaceae, Marchantiophyta) species from the deep south of Aotearoa-New Zealand based on an integrated evidence-based approach. PhytoKeys 8: 1336. https://doi.org/10.3897/phytokeys.8.2496" title="Frullania knightbridgei, a new liverwort (Frullaniaceae, Marchantiophyta) species from the deep south of Aotearoa-New Zealand based on an integrated evidence-based approach." volume="8" year="2012">2012</bibRefCitation>
,
<bibRefCitation id="6291D524FBFF58982817AFB0ED21617B" DOI="10.2478/pbj-2013-0056" author="von Konrat" firstAuthor="von Konrat" journalOrPublisher="Polish Botanical Journal" pagination="437-447" refId="B139" refString="von Konrat M, de Lange P, Larraín J, Hentschel J, Carter B, Shaw J, Shaw B (2013) A small world: uncovering hidden diversity in Frullania a new species from Aotearoa-New Zealand. Polish Botanical Journal 58: 437447. https://doi.org/10.2478/pbj-2013-0056" title="A small world: uncovering hidden diversity in Frullania - a new species from Aotearoa-New Zealand." volume="58" year="2013">2013</bibRefCitation>
;
<bibRefCitation id="46FAAF8EB7D9E9EF9AF9A0BB13D6BD1C" DOI="10.1016/j.ympev.2016.10.002" author="Carter" etAl="et al." firstAuthor="Carter" journalOrPublisher="Molecular Phylogenetics and Evolution" pagination="16-26" refId="B10" refString="Carter BE, Larraín J, Manukjanová A, Shaw B, Shaw AJ, Heinrichs J, de Lange P, Suleiman M, Thouvenot L, von Konrat M (2017) Species delimitation and biogeography of a southern hemisphere liverwort clade, Frullania subgenus Microfrullania (Frullaniaceae, Marchantiophyta). Molecular Phylogenetics and Evolution 107: 1626. https://doi.org/10.1016/j.ympev.2016.10.002" title="Species delimitation and biogeography of a southern hemisphere liverwort clade, Frullania subgenus Microfrullania (Frullaniaceae, Marchantiophyta)." volume="107" year="2017">Carter et al. 2017</bibRefCitation>
;
<bibRefCitation id="F06B46CC4F272E2F8D5BE5651D497125" DOI="10.1080/14772000.2016.1217096" author="Silva" etAl="et al." firstAuthor="Silva" journalOrPublisher="Systematics and Biodiversity" pagination="156-165" refId="B122" refString="Silva SE, Silva DN, Almeida T, Garcia CA, Paulo OS, Sim-Sim M (2017) Age estimates of Frullania (Frullaniaceae, Porellales) main lineages: another example of rapid and recent diversification in liverwort evolution. Systematics and Biodiversity 15: 156165. https://doi.org/10.1080/14772000.2016.1217096" title="Age estimates of Frullania (Frullaniaceae, Porellales) main lineages: another example of rapid and recent diversification in liverwort evolution." volume="15" year="2017">Silva et al. 2017</bibRefCitation>
;
<bibRefCitation id="D00FDE2A05F83584948522F1B46888DB" DOI="10.1127/nova-suppl/2020/201" author="Mamontov" etAl="et al." firstAuthor="Mamontov" journalOrPublisher="Nova Hedwigia Beiheft" pagination="201-242" refId="B88" refString="Mamontov YS, Vilnet AA, Atwood JJ, Konstantinova NA (2020) Molecular phylogenetic study of Frullania subsect. Inflatae (Frullaniaceae, Marchantiophyta) in the Holarctic with description of a new subgenus and three new species. Nova Hedwigia Beiheft 150: 201242. https://doi.org/10.1127/nova-suppl/2020/201" title="Molecular phylogenetic study of Frullania subsect. Inflatae (Frullaniaceae, Marchantiophyta) in the Holarctic with description of a new subgenus and three new species." volume="150" year="2020">Mamontov et al. 2020</bibRefCitation>
;
<bibRefCitation id="B45BEC42D5D4578ECC32E092A71FFBC4" author="Schafer-Verwimp and Winter" firstAuthor="Schafer-Verwimp" journalOrPublisher="Frahmia" pagination="1-14" refId="B112" refString="Schäfer-Verwimp A, Winter G (2020) Frullania dasyueshanensis, a new species with terete perianth from Taiwan (Marchantiophyta: Frullaniaceae). Frahmia 20: 114. http://www.frahmia.de/downloads/frahmia/frahmia_0020.pdf" title="Frullania dasyueshanensis, a new species with terete perianth from Taiwan (Marchantiophyta: Frullaniaceae)." url="http://www.frahmia.de/downloads/frahmia/frahmia_0020.pdf" volume="20" year="2020">Schäfer-Verwimp and Winter 2020</bibRefCitation>
,
<bibRefCitation id="F4F40DBE3107484FD25996B3665D031C" author="Schafer-Verwimp and Winter" firstAuthor="Schafer-Verwimp" journalOrPublisher="Frahmia" pagination="1-12" refId="B113" refString="Schäfer-Verwimp A, Winter G (2022) Frullania azureomontana a new species from Jamaica (Marchantiophyta: Frullaniaceae). Frahmia 30: 112. http://www.frahmia.de/downloads/frahmia/frahmia_0030.pdf" title="Frullania azureomontana - a new species from Jamaica (Marchantiophyta: Frullaniaceae)." url="http://www.frahmia.de/downloads/frahmia/frahmia_0030.pdf" volume="30" year="2022">2022</bibRefCitation>
,
<bibRefCitation id="01879776453386767E1B2A7DEFEA1C62" author="Schafer-Verwimp and Winter" firstAuthor="Schafer-Verwimp" journalOrPublisher="Frahmia" pagination="1-14" refId="B114" refString="Schäfer-Verwimp A, Winter G (2023) Frullania hohenesteri (Marchantiophyta: Frullaniaceae) a new species from Santa Catarina, South Brazil. Frahmia 39: 114. http://www.frahmia.de/downloads/frahmia/frahmia_0039.pdf" title="Frullania hohenesteri (Marchantiophyta: Frullaniaceae) - a new species from Santa Catarina, South Brazil." url="http://www.frahmia.de/downloads/frahmia/frahmia_0039.pdf" volume="39" year="2023">2023</bibRefCitation>
). Consequently, the assessment of morphological differences between
<taxonomicName id="41CBFDDF1BA483D1597A3F2B2F10EA52" authorityName="Mamontov, K. Feldberg, Schaf. - Verw., Gradst." authorityYear="2025" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="chiapasensis">
<emphasis id="F39DA571F7E5924197B067170D023BBF" italics="true">F. chiapasensis</emphasis>
</taxonomicName>
and the extant species of the genus is difficult. Moreover, the morphological homoplasies within
<taxonomicName id="4A9679219DD055DA1683668024B3327F" authorityName="Raddi" authorityYear="1818" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="genus">
<emphasis id="E62637C0EB3740AFD93AD4EEFBB4670D" italics="true">Frullania</emphasis>
</taxonomicName>
challenge the subgeneric assignment of numerous extant non-sequenced species, and even more so that of fossils. Based on the combination of the characteristics of leaf lobes, lobules, and underleaves,
<taxonomicName id="2CF1EB1FF7F48B61E79F55A794697C02" authorityName="Mamontov, K. Feldberg, Schaf. - Verw., Gradst." authorityYear="2025" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="chiapasensis">
<emphasis id="10435EE7900CDF063B112AAC0D0690A3" italics="true">F. chiapasensis</emphasis>
</taxonomicName>
is similar to species of several subgenera: namely
<taxonomicName id="939F009DED1411BC286A0787EB7A572F" class="Jungermanniopsida" family="Frullaniaceae" genus="Caulisequa" higherTaxonomySource="CoL" kingdom="Plantae" order="Frullaniales" phylum="Marchantiophyta" rank="genus">
<emphasis id="C0CC847A1492D55E4122168B2087122C" italics="true">Caulisequa</emphasis>
</taxonomicName>
,
<emphasis id="25678255DAE0CAEF6D5EEAB974A55E35" italics="true">Diastaloba</emphasis>
I,
<emphasis id="3FAA167428C53EED2DE32AED31A53877" italics="true">Diastaloba</emphasis>
II,
<emphasis id="98D6C84C98E7DEE8EE5D706AD1E083E3" italics="true">Diastaloba</emphasis>
III,
<taxonomicName id="E2ADE79F1E2684358EC42E737E1309FF" authority=", Mammillosae" authorityName="Mammillosae" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="genus">
<emphasis id="CBF8F8E373514E9B88BCB7908EE4AA3F" italics="true">Frullania</emphasis>
,
<emphasis id="F9A869940006DE09CAA375191DF89068" italics="true">Mammillosae</emphasis>
</taxonomicName>
,
<taxonomicName id="C6C1115D68AF91D213DF7010BD212830" class="Bryopsida" family="Meteoriaceae" genus="Meteoriopsis" kingdom="Plantae" order="Leucodontales" phylum="Bryophyta" rank="genus">
<emphasis id="F9FC0DB31D08FB3EF1A4392914C5EA9C" italics="true">Meteoriopsis</emphasis>
</taxonomicName>
, and
<taxonomicName id="C68BD12EBDF3EA091876A0E464F6ED08" rank="subGenus" subGenus="Microfrullania">
<emphasis id="B5654F6D5DB43B717559E37FBCD85FEA" italics="true">Microfrullania</emphasis>
</taxonomicName>
. To compare them with
<taxonomicName id="CA0E027F9EF6FCD004D40305B54A2819" authorityName="Mamontov, K. Feldberg, Schaf. - Verw., Gradst." authorityYear="2025" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="chiapasensis">
<emphasis id="F880FA9E9DCCA26EAB15552F1CF4F416" italics="true">F. chiapasensis</emphasis>
</taxonomicName>
, we have analyzed the information (protologues, descriptions, and figures) for all species of the mentioned subgenera, which are accepted in
<bibRefCitation id="EEF5833BB22D539270B3004319783973" DOI="10.3897/phytokeys.59.6261" author="Söderström L &amp; Hagborg A &amp; von Konrat M &amp; Bartholomew-Began S &amp; Bell D &amp; Briscoe L &amp; Brown E &amp; Cargill DC &amp; Costa DP &amp; Crandall-Stotler BJ &amp; Cooper ED &amp; Dauphin G &amp; Engel JJ &amp; Feldberg K &amp; Glenny D &amp; Gradstein SR &amp; He X &amp; Heinrichs J &amp; Hentschel J &amp; Ilkiu-Borges AL &amp; Katagiri T &amp; Konstantinova NA &amp; Larraín J &amp; Long DG &amp; Nebel M &amp; Pócs T &amp; Puche F &amp; Reiner-Drehwald E &amp; Renner MAM &amp; Sass-Gyarmati A &amp; Schäfer-Verwimp A &amp; Moragues JGS &amp; Stotler RE &amp; Sukkharak P &amp; Thiers BM &amp; Uribe J &amp; Váňa J &amp; Villarreal JC &amp; Wigginton M &amp; Zhang L &amp; Zhu R-L" etAl="et al." firstAuthor="Soderstrom" journalOrPublisher="PhytoKeys" pagination="1-828" refId="B123" refString="Söderström L, Hagborg A, von Konrat M, Bartholomew-Began S, Bell D, Briscoe L, Brown E, Cargill DC, Costa DP, Crandall-Stotler BJ, Cooper ED, Dauphin G, Engel JJ, Feldberg K, Glenny D, Gradstein SR, He X, Heinrichs J, Hentschel J, Ilkiu-Borges AL, Katagiri T, Konstantinova NA, Larraín J, Long DG, Nebel M, Pócs T, Puche F, Reiner-Drehwald E, Renner MAM, Sass-Gyarmati A, Schäfer-Verwimp A, Moragues JGS, Stotler RE, Sukkharak P, Thiers BM, Uribe J, Váňa J, Villarreal JC, Wigginton M, Zhang L, Zhu R-L (2016) World checklist of hornworts and liverworts. PhytoKeys 59: 1828. https://doi.org/10.3897/phytokeys.59.6261" title="World checklist of hornworts and liverworts." volume="59" year="2016">Söderström et al. (2016)</bibRefCitation>
. Some members of these subgenera are well distinguished from
<taxonomicName id="C3DDC84543E38474624E4058DEE71872" authorityName="Mamontov, K. Feldberg, Schaf. - Verw., Gradst." authorityYear="2025" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="chiapasensis">
<emphasis id="4179C3EA1E5D8278F0DD44E62742FC22" italics="true">F. chiapasensis</emphasis>
</taxonomicName>
by the presence of moniliate ocelli in leaf lobes or by the shape of the underleaves (if the underleaves are entire at apex or appendiculate at base, for example). However, many taxa can be distinguished from the newly described species only with difficulty. Therefore, we compare extant species similar to
<taxonomicName id="5B0B7CABD3C87533CF924620161C6809" authorityName="Mamontov, K. Feldberg, Schaf. - Verw., Gradst." authorityYear="2025" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="chiapasensis">
<emphasis id="94262E393129EE124D266E73F2658429" italics="true">F. chiapasensis</emphasis>
</taxonomicName>
in a morphology matrix based on 5 + 11 critical characters (see material &amp; methods and Appendix 1). Most species of subgenera
<taxonomicName id="A5D746F4F9D62D7FAE4E592771F10A91" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="genus">
<emphasis id="E0127626A52C3C72CAEAB5ADD5EEE907" italics="true">Frullania</emphasis>
</taxonomicName>
and
<taxonomicName id="08570E0B8C35BC43259750E9BE9DEA6B" class="Bryopsida" family="Meteoriaceae" genus="Meteoriopsis" kingdom="Plantae" order="Leucodontales" phylum="Bryophyta" rank="genus">
<emphasis id="700B3EBA7228E14F0950FFDB49CCAD2E" italics="true">Meteoriopsis</emphasis>
</taxonomicName>
were excluded from consideration based on characters 15, while
<taxonomicName id="7956F04A262570290DAF21A1EEB4EC85" authorityName="Mamontov, K. Feldberg, Schaf. - Verw., Gradst." authorityYear="2025" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="chiapasensis">
<emphasis id="E1382EB465731BE1792F1D348D263BF8" italics="true">F. chiapasensis</emphasis>
</taxonomicName>
+ 144 species meeting these criteria have been chosen to map their approximate differences and similarities based on 11 additional characters.
</paragraph>
<paragraph id="ACD68EA23AF527D12707999EDCC36865">
Based on the character comparison, the extant species morphologically most similar to
<taxonomicName id="01C325515B3D4BAC3CEE45FD1F6B9ED0" authorityName="Mamontov, K. Feldberg, Schaf. - Verw., Gradst." authorityYear="2025" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="chiapasensis">
<emphasis id="3483530271E1918B341FB37291D49149" italics="true">F. chiapasensis</emphasis>
</taxonomicName>
is
<taxonomicName id="47226E1D8E5AC0556B9B2BC82485A757" authority="Steph." authorityName="Steph." class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="simmondsii">
<emphasis id="6CFB60F78D8A92D4C0C5CE3BC007DC1E" italics="true">F. simmondsii</emphasis>
Steph.
</taxonomicName>
, as described in
<bibRefCitation id="03B3B287D3F00BA41B3B237A55DBBAE6" author="Hattori" firstAuthor="Hattori" journalOrPublisher="Journal of the Hattori Botanical Laboratory" pagination="119-153" refId="B53" refString="Hattori S (1979) A revision of the Australasian species of the genus Frullania, Hepaticae, II. Journal of the Hattori Botanical Laboratory 46: 119153. https://www.jstage.jst.go.jp/article/jhbl/46/0/46_119/_pdf" title="A revision of the Australasian species of the genus Frullania, Hepaticae, II." url="https://www.jstage.jst.go.jp/article/jhbl/46/0/46_119/_pdf" volume="46" year="1979">Hattori (1979</bibRefCitation>
, fig. 42). According to Appendix 1, the species coincide with each other in all 11 characters. However, based on the description and illustration in
<bibRefCitation id="757FBCD8DED35C488F689E9D3BF8D132" author="Hattori S" firstAuthor="Hattori" journalOrPublisher="Journal of the Hattori Botanical Laboratory" pagination="119-153" refId="B53" refString="Hattori S (1979) A revision of the Australasian species of the genus Frullania, Hepaticae, II. Journal of the Hattori Botanical Laboratory 46: 119153. https://www.jstage.jst.go.jp/article/jhbl/46/0/46_119/_pdf" title="A revision of the Australasian species of the genus Frullania, Hepaticae, II." url="https://www.jstage.jst.go.jp/article/jhbl/46/0/46_119/_pdf" volume="46" year="1979">Hattori (1979)</bibRefCitation>
, the stylus of
<taxonomicName id="E000FC4BDD087E660731E7558391864F" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="simmondsii">
<emphasis id="96C54928FA742416B92FB004D0D4EBA6" italics="true">F. simmondsii</emphasis>
</taxonomicName>
is similar to that in numerous species of subg.
<emphasis id="793C5E9A80404FC95B9DE4346362A4C0" italics="true">Diastaloba</emphasis>
s. l. by leaving a narrow foliose strip extending along the keel, and thus being wider than long. In contrast, the stylus of
<taxonomicName id="E94241D1BB3B775D95E5E915FC8DBA10" authorityName="Mamontov, K. Feldberg, Schaf. - Verw., Gradst." authorityYear="2025" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="chiapasensis">
<emphasis id="01C004F3ECF9D8F461CC86F65F15E497" italics="true">F. chiapasensis</emphasis>
</taxonomicName>
resembles the foliaceous styli found in numerous species of subg.
<emphasis id="6E62CD8D76C141162FFA89CFFE7E9D8C" italics="true">Trachycolea</emphasis>
by being longer than wide. Summarizing all differences between both species,
<taxonomicName id="64EA90DEBF69EE719C316D279BA28863" authorityName="Mamontov, K. Feldberg, Schaf. - Verw., Gradst." authorityYear="2025" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="chiapasensis">
<emphasis id="CD2869347B8BA0C3FC0FD64E73A9E25D" italics="true">F. chiapasensis</emphasis>
</taxonomicName>
can be distinguished from
<taxonomicName id="E2F50EE191F10E01FE6ACCD9971CE0CD" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="simmondsii">
<emphasis id="82490DD490F819E317EF4B620523E9C5" italics="true">F. simmondsii</emphasis>
</taxonomicName>
as follows: (1) leaf lobes longer than wide in
<taxonomicName id="0F8AC6A20A80AD492D79621BBAE8EC22" authorityName="Mamontov, K. Feldberg, Schaf. - Verw., Gradst." authorityYear="2025" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="chiapasensis">
<emphasis id="588555F7F38F2C8FEDEE26289E98AF03" italics="true">F. chiapasensis</emphasis>
</taxonomicName>
vs. wider than long in
<taxonomicName id="FE68278FC257453B7058C4856CFA139E" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="simmondsii">
<emphasis id="96AF376A9FDA46D7D7D43913F639A985" italics="true">F. simmondsii</emphasis>
</taxonomicName>
; (2) stylus foliaceous, longer than wide vs. filiform, when flat narrowly foliaceous, wider than long; (3) stem underleaves mostly longer than wide, narrowly obcuneate at base, with the margins concave in the lower two-thirds vs. stem underleaves mostly wider than long, widely obcuneate at base, with the margins largely convex in lower two-thirds.
</paragraph>
<paragraph id="86CACE16524C1C03F2A41BDB5A2B8B14">
Besides
<taxonomicName id="7160B0CD6431FBA1A53B4084CD6D6670" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="simmondsii">
<emphasis id="D0615E5A8EDEA3AEBD38192A54EA2865" italics="true">F. simmondsii</emphasis>
</taxonomicName>
, we have identified eight more species, each differing from
<taxonomicName id="46ED47B67A9AB2169DCB5F82477A130C" authorityName="Mamontov, K. Feldberg, Schaf. - Verw., Gradst." authorityYear="2025" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="chiapasensis">
<emphasis id="163FAFED414A70E32A09FDCE80A9157D" italics="true">F. chiapasensis</emphasis>
</taxonomicName>
in one of the listed 11 characters (Appendix 1). The species are
<taxonomicName id="49A31407751A738AF25ACFC73DD9ACF5" authority="von Konrat et al." authorityName="von Konrat" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="colliculosa">
<emphasis id="AE6A8BD53C2637F17873021F1E9994D2" italics="true">F. colliculosa</emphasis>
von Konrat et al.
</taxonomicName>
,
<taxonomicName id="62F06F0C48387CA2EFF7E129961E7CF2" authority="Steph." authorityName="Steph." class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="eplicata">
<emphasis id="9841BEAD1C50A87B7473CDA8578E26B2" italics="true">F. eplicata</emphasis>
Steph.
</taxonomicName>
,
<taxonomicName id="3FD564DA8390E2A8F8B4CC3C488150AF" authority="Vanden Berghen" authorityName="Vanden Berghen" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="gabonensis">
<emphasis id="A647FDAC2E6EC3B3A318B8B35C742213" italics="true">F. gabonensis</emphasis>
Vanden Berghen
</taxonomicName>
,
<taxonomicName id="FD322484BFE43C117F014B4C6BE22CB8" authority="von Konrat et al." authorityName="von Konrat" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="hodgsoniae">
<emphasis id="E07FE7FBC814751A2BCA50BA8D7DD8DA" italics="true">F. hodgsoniae</emphasis>
von Konrat et al.
</taxonomicName>
,
<taxonomicName id="A29B7F7B1E6E8B972CFCA0EAEC98E7EF" authority="S. Hatt." authorityName="S. Hatt." class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="multilaceroides">
<emphasis id="343451C6FC7C96AFC47278BDF1AA5B24" italics="true">F. multilaceroides</emphasis>
S. Hatt.
</taxonomicName>
,
<taxonomicName id="8FE42037B822AA8071FA2E678EAC76A3" authority="S. Hatt." authorityName="S. Hatt." class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="scalaris">
<emphasis id="7AFA046E43A206A03FB08426119ACBC8" italics="true">F. scalaris</emphasis>
S. Hatt.
</taxonomicName>
,
<taxonomicName id="511C17F52EF643A6C43997F1FD557059" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="subtilissima">
<emphasis id="9CFA974AEEC70E2412EE84F18192B164" italics="true">F. subtilissima</emphasis>
</taxonomicName>
(=
<taxonomicName id="C4B9FCA1A51539021CBDEB6BC3908086" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="taxodiocola">
<emphasis id="FAB96127ABCA9F3E7B88EFA76BAA2F9C" italics="true">F. taxodiocola</emphasis>
</taxonomicName>
and
<taxonomicName id="8D8BCE52A87F407F1D9A40402C13BC51" authority="Taylor" authorityName="Taylor" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="exilis">
<emphasis id="B2049F5BCABF5666B5F773BC964D963C" italics="true">F. exilis</emphasis>
Taylor
</taxonomicName>
) and
<taxonomicName id="59080988E5BCD2B96E8CC6F9537118F2" authority="Mitt." authorityName="Mitt." class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="vaga">
<emphasis id="3E139C3E381857FF68657800BB301A3D" italics="true">F. vaga</emphasis>
Mitt.
</taxonomicName>
Among them, five species (
<taxonomicName id="AD7D8C61F961CDAB40AB9FF67F308D74" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="colliculosa">
<emphasis id="0287184E27D683D7673F75A899084E66" italics="true">F. colliculosa</emphasis>
</taxonomicName>
,
<taxonomicName id="2715EFA90367690B4775DCF2DBF8F0C5" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="gabonensis">
<emphasis id="1B8308C016BFB6C0B3900FB7A690C061" italics="true">F. gabonensis</emphasis>
</taxonomicName>
,
<taxonomicName id="3B9111BBA233244C8F476311E7D7E31B" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="hodgsoniae">
<emphasis id="0A2C9B0225057A6E4D5680089D093110" italics="true">F. hodgsoniae</emphasis>
</taxonomicName>
,
<taxonomicName id="86EF6F31F98CF4B9D8FE6B29C746CB9C" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="multilaceroides">
<emphasis id="F3E2099B6F646C6716E8BB6B8337E806" italics="true">F. multilaceroides</emphasis>
</taxonomicName>
,
<taxonomicName id="44D545981663AEB257D20E89768C0873" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="subtilissima">
<emphasis id="F06AEBDA670FCE4F220DB2D0FEED542F" italics="true">F. subtilissima</emphasis>
</taxonomicName>
) differ from
<taxonomicName id="ABDE72B0CC0EEB41695CD45195FDE7B3" authorityName="Mamontov, K. Feldberg, Schaf. - Verw., Gradst." authorityYear="2025" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="chiapasensis">
<emphasis id="D4BD5E14290C3BD64E515F380F97C940" italics="true">F. chiapasensis</emphasis>
</taxonomicName>
in the lobule arrangement. The lobules in the mentioned five species are obliquely patent, spreading from the stem at angles of 2555 ° or more, whereas in
<taxonomicName id="96F7ECE9A91B316DF4165070AE6F280C" authorityName="Mamontov, K. Feldberg, Schaf. - Verw., Gradst." authorityYear="2025" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="chiapasensis">
<emphasis id="51623E2DF5127F595E58D088E7EFE270" italics="true">F. chiapasensis</emphasis>
</taxonomicName>
the lobules are slightly patent or largely parallel to the stem. Moreover, in
<taxonomicName id="EF47172E83BD381BDEE464841E08F576" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="colliculosa">
<emphasis id="FF1FE2DB89D3CB93AD3288E9BD43549C" italics="true">F. colliculosa</emphasis>
</taxonomicName>
,
<taxonomicName id="2BF9D9063038C4D8D7059DA51B8B2CEE" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="gabonensis">
<emphasis id="DEEF2AB948123BB9EA610D53C9AE1CF1" italics="true">F. gabonensis</emphasis>
</taxonomicName>
,
<taxonomicName id="A63BE22D05D2F8447A63136641AFC8A2" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="hodgsoniae">
<emphasis id="6D09FB2835CAF68D62EAB65F1D88F248" italics="true">F. hodgsoniae</emphasis>
</taxonomicName>
, and
<taxonomicName id="FD33FFAD65A9CF28C192216441512E66" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="multilaceroides">
<emphasis id="C508B56CCDCEF4360769A03A4EBF51CA" italics="true">F. multilaceroides</emphasis>
</taxonomicName>
the underleaves of leading stems are usually slightly broader than long and broadest in the middle, whereas in
<taxonomicName id="D050411E13533FD474191FFABD868CEE" authorityName="Mamontov, K. Feldberg, Schaf. - Verw., Gradst." authorityYear="2025" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="chiapasensis">
<emphasis id="E13C671F08ED3E70EB245BDBC72687F2" italics="true">F. chiapasensis</emphasis>
</taxonomicName>
the underleaves are mostly longer than wide and broadest at the upper third. In the studied extant specimens of
<taxonomicName id="2481DC513B5A434234EFF0C515C5B097" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="subtilissima">
<emphasis id="54FD8B7F842BB9851C5C702CFC92B1B1" italics="true">F. subtilissima</emphasis>
</taxonomicName>
from Costa-Rica (Schäfer-Verwimp &amp; Holz SV / H- 0486 / A, Schäfer-Verwimp &amp; Holz SV / H- 0429 / Z,
<named-content id="FF04D3DDEAF06D496E9EB2F965D5BDE0" content-type="dwc:institutional_code" xlink_href="http://grbio.org/institution/main-botanical-garden-russian-academy-sciences" xlink_title="Russian Academy of Sciences">MHA</named-content>
), the width and length of leaf lobes exceed the length of leaf lobules ca. 2 and 2.8 times, respectively, thus the leaf lobes are ca. 810 times larger than lobules, while the lobe apex is often apiculate and the margins of underleaves entire or barely angulate. In contrast, in
<taxonomicName id="D8B19B127916E6A3B68350010ADA707F" authorityName="Mamontov, K. Feldberg, Schaf. - Verw., Gradst." authorityYear="2025" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="chiapasensis">
<emphasis id="1948A4309C57BAA032126BBAC3F50D46" italics="true">F. chiapasensis</emphasis>
</taxonomicName>
the length and width of leaf lobes exceeds the length of leaf lobules ca. 1.4 and 1.6 times, respectively, thus the leaf lobes are only ca. 23 times larger than lobules, while the lobe apex is obtuse to rounded, and the margins of underleaves are armed with 12 teeth of 13 cells on both sides. Among the remaining three species,
<taxonomicName id="EAB2191BEF3FFC2C5E5ABA145849389B" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="eplicata">
<emphasis id="F641DDB254F8680C6A592A04AB3EF18B" italics="true">F. eplicata</emphasis>
</taxonomicName>
differs from
<taxonomicName id="CC1E1E605497A2DF840A80BDCD1AC070" authorityName="Mamontov, K. Feldberg, Schaf. - Verw., Gradst." authorityYear="2025" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="chiapasensis">
<emphasis id="15936ED2E62019337C260FF8C02DEECC" italics="true">F. chiapasensis</emphasis>
</taxonomicName>
by the low position of the lobule mouth apex that is similar to species of subgenera
<taxonomicName id="F1CE537D41658B6957E0DBACEFF5E003" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="genus">
<emphasis id="560C5B9B253830FBE9EE7781F5375EDB" italics="true">Frullania</emphasis>
</taxonomicName>
,
<taxonomicName id="230065442D48476EE82863AEDB98DBE5" class="Bryopsida" family="Meteoriaceae" genus="Meteoriopsis" kingdom="Plantae" order="Leucodontales" phylum="Bryophyta" rank="genus">
<emphasis id="BF4EABFAA321D467FA0EF0B50D313CF3" italics="true">Meteoriopsis</emphasis>
</taxonomicName>
, and
<emphasis id="E406FEC5196095E2ACA4EBD26B628361" italics="true">Trachycolea</emphasis>
. Moreover, well-developed underleaves in
<taxonomicName id="FD943CA83AAE4E7A59FB307922BEA7C1" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="eplicata">
<emphasis id="ADDCA8CEDF3F512A03DD039FCB5E4547" italics="true">F. eplicata</emphasis>
</taxonomicName>
are up to 3 times broader than the stem and wider than long (Vanden Bergen 1976, fig. 21 A). The second species,
<taxonomicName id="5DCBBC7F58B75BEB764F4AC837EC45C7" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="scalaris">
<emphasis id="26A8A73D44E870004F66B99D0AA2D089" italics="true">F. scalaris</emphasis>
</taxonomicName>
, is very similar to
<taxonomicName id="5FAAF48772CB474B70B9A8FAF3802D66" authorityName="Mamontov, K. Feldberg, Schaf. - Verw., Gradst." authorityYear="2025" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="chiapasensis">
<emphasis id="1684DD90C76C777E2C6B2EAE70F46F47" italics="true">F. chiapasensis</emphasis>
</taxonomicName>
in the shape and size of leaf lobes and lobules. However, the leaf lobules in the former species are apically overlapping the stem, while the underleaves are described and illustrated as only slightly broader than the stem, entire margined, broadest in the middle, almost bilobed to 0.5 of their length, with triangular-lanceolate lobes (
<bibRefCitation id="C29D2BF3E683C5858F2C61A16D6EF2C4" author="Hattori" firstAuthor="Hattori" journalOrPublisher="Journal of the Hattori Botanical Laboratory" pagination="409-438" refId="B52" refString="Hattori S (1977) Dr. H. Hürlimanns collection of New Caledonian Frullaniaceae. Journal of the Hattori Botanical Laboratory 43: 409438. https://www.jstage.jst.go.jp/article/jhbl/43/0/43_409/_pdf/-char/ja" title="Dr. H. Hürlimanns collection of New Caledonian Frullaniaceae." url="https://www.jstage.jst.go.jp/article/jhbl/43/0/43_409/_pdf/-char/ja" volume="43" year="1977">Hattori 1977</bibRefCitation>
, fig. 15). In contrast, in
<taxonomicName id="688F3CF3268D8FEE5EBE129BCC057626" authorityName="Mamontov, K. Feldberg, Schaf. - Verw., Gradst." authorityYear="2025" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="chiapasensis">
<emphasis id="7AED84ABFA068B8FF6F6149807EB9261" italics="true">F. chiapasensis</emphasis>
</taxonomicName>
the leaf lobules are mostly parallel to the stem, while the underleaves are 2.12.5 times broader than stem, toothed in the upper third, bilobed to 0.20.3 of their length, with widely triangular lobes. The last species,
<taxonomicName id="58E234B0AAF31DA9FDB4ED3BF0FBEC40" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="vaga">
<emphasis id="8DE1025A30C8A3C2312AC2C7FCF9CA82" italics="true">F. vaga</emphasis>
</taxonomicName>
, is described to have only shallowly bilobed, entire margined underleaves (
<bibRefCitation id="7DFF2340B644C413D155D07F83296580" DOI="10.5962/bhl.title.455" author="Mitten" editor="Seemann B" firstAuthor="Mitten" journalOrPublisher="2. Reeve, London" pagination="404-418" refId="B93" refString="Mitten W (1865) Jungermanniae. In: Seemann B (Ed.) Flora Vitiensis, v. 2. Reeve, London, 404418. https://doi.org/10.5962/bhl.title.455" title="Jungermanniae." volumeTitle="Flora Vitiensis, v." year="1865">Mitten 1865</bibRefCitation>
), while other distinctions are not evident from the protologue.
</paragraph>
<paragraph id="85FA9115BC58AC3C85355616062F9F6D">
The remaining 135 species in Appendix 1, apart from the 9 species discussed above, can be distinguished from
<taxonomicName id="693E93135A454EC5AC45100ED99411CB" authorityName="Mamontov, K. Feldberg, Schaf. - Verw., Gradst." authorityYear="2025" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="chiapasensis">
<emphasis id="027F4891DBC3B2BAF51D2FDF8A86EDD4" italics="true">F. chiapasensis</emphasis>
</taxonomicName>
by two to seven characters. Therefore, the latter taxon can be considered as a separate species which essentially differs from all the described extant and fossil
<taxonomicName id="0AE2BF7D5B5CBCBD1EFEC1F9EFC861BA" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="undetermined">
<emphasis id="AA5FC95985B8659756272CCBC0EF028E" italics="true">Frullania</emphasis>
species.
</taxonomicName>
The subgeneric assignment of
<taxonomicName id="1655564286E47F735D2696A4596734B8" authorityName="Mamontov, K. Feldberg, Schaf. - Verw., Gradst." authorityYear="2025" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="chiapasensis">
<emphasis id="6813AF6A08730F2E7B22223D28BD636C" italics="true">F. chiapasensis</emphasis>
</taxonomicName>
remains problematic because the species morphologically most similar to it are treated now as belonging to different subgenera, namely
<emphasis id="7B4BFA96359DE278844250570D6D03BB" italics="true">Diastaloba</emphasis>
I and
<taxonomicName id="C796D307363C49DA309690BAA152BEFE" rank="subGenus" subGenus="Microfrullania">
<emphasis id="43683D7FC4DE498761019892A3F9D5C7" italics="true">Microfrullania</emphasis>
</taxonomicName>
(
<bibRefCitation id="16F7786140879DBD9FF0B6F172B7717A" DOI="10.3897/phytokeys.59.6261" author="Soderstrom" etAl="et al." firstAuthor="Soderstrom" journalOrPublisher="PhytoKeys" pagination="1-828" refId="B123" refString="Söderström L, Hagborg A, von Konrat M, Bartholomew-Began S, Bell D, Briscoe L, Brown E, Cargill DC, Costa DP, Crandall-Stotler BJ, Cooper ED, Dauphin G, Engel JJ, Feldberg K, Glenny D, Gradstein SR, He X, Heinrichs J, Hentschel J, Ilkiu-Borges AL, Katagiri T, Konstantinova NA, Larraín J, Long DG, Nebel M, Pócs T, Puche F, Reiner-Drehwald E, Renner MAM, Sass-Gyarmati A, Schäfer-Verwimp A, Moragues JGS, Stotler RE, Sukkharak P, Thiers BM, Uribe J, Váňa J, Villarreal JC, Wigginton M, Zhang L, Zhu R-L (2016) World checklist of hornworts and liverworts. PhytoKeys 59: 1828. https://doi.org/10.3897/phytokeys.59.6261" title="World checklist of hornworts and liverworts." volume="59" year="2016">Söderström et al. 2016</bibRefCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>

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@ -0,0 +1,575 @@
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<mods:affiliation id="106DB8222917967C2B4484DA050220EE">Finnish Museum of Natural History, University of Helsinki, P. O. Box 7, 00014 Helsinki, Finland</mods:affiliation>
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<treatment id="B8AF380D312455309F95494016EBD03E" ID-arpha="B8AF380D-3124-5530-9F95-494016EBD03E" ID-zoobank="https://zoobank.org/DAB13777-0E72-4E4A-9CA3-07E59DF17506" LSID="urn:lsid:plazi:treatment:B8AF380D312455309F95494016EBD03E" LSID-ZBK="urn:lsid:zoobank.org:act:DAB13777-0E72-4E4A-9CA3-07E59DF17506" httpUri="http://treatment.plazi.org/id/B8AF380D312455309F95494016EBD03E" scope_class="Jungermanniopsida" scope_family="Lejeuneaceae" scope_order="Porellales" scope_phylum="Marchantiophyta" scope_subClass="Jungermanniidae" scope_subFamily="Ptychanthoideae">
<subSubSection id="317914D598F9BA7D970B536CFAE94FFB" type="nomenclature">
<paragraph id="F974ECA63D28AC23CF3B5919A27EA96A">
<taxonomicName id="E8BEBF6B7B96F77F9874CAD64FA7024B" authority="K. Feldberg, Gradst., Schaf.-Verw., Mamontov" authorityName="K. Feldberg, Gradst., Schaf. - Verw., Mamontov" authorityYear="2025" class="Jungermanniopsida" family="Lejeuneaceae" genus="Thysananthus" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="patrickmuelleri" status="sp. nov.">
<emphasis id="045C7CDD7C9B96AACA0405A1AD0BD752" italics="true">Thysananthus patrickmuelleri</emphasis>
K. Feldberg, Gradst., Schäf.-Verw., Mamontov
</taxonomicName>
<taxonomicNameLabel id="D3EC91C728AA8AE5DE3C8FE5ED4E8467" rank="species">sp. nov.</taxonomicNameLabel>
</paragraph>
</subSubSection>
<subSubSection id="SECID0EL2BG" type="Holotype">
<paragraph id="C80AAC94DD5618F319EF629D9EED170C">
<heading id="C8010EB1D3644D9FF692FF74FD4559DE" reason="title">
<typeStatus id="BD4CB5987EE3746D83D0E8CAE5D8ECB0" type="holotype">Holotype</typeStatus>
.
</heading>
</paragraph>
<paragraph id="6BD684544FDEC8C876B29D3FB82CCB66">
<materialsCitation id="61977E63317D41412A78761FAE6F8DCD" collectionCode="GZG" specimenCode="GZG.BST.22086" specimenCount="1" typeStatus="holotype">
Geoscientific Collection of the University of Göttingen, Germany (
<abbrev content-type="institution" id="ABBRID0ER2BG" xlink_title="Geoscientific Collection of the University of Göttingen">
<collectionCode id="3DADD7E735803F4A1E8F7239EEB3F175">GZG</collectionCode>
</abbrev>
),
<specimenCode id="B34A2B0C1ADD6430ED86F7A379A76A94">
<abbrev content-type="institution" id="ABBRID0EW2BG" xlink_title="Geoscientific Collection of the University of Göttingen">GZG</abbrev>
.BST.22086
</specimenCode>
.
</materialsCitation>
</paragraph>
</subSubSection>
<subSubSection id="SECID0E22BG" type="material">
<paragraph id="06394560A060EE3644CF55597D045993">
<heading id="8195FED211004707F9B4A5B3829E9B4C" reason="title">Additional specimens examined.</heading>
</paragraph>
<paragraph id="316F9AF9D3F52B46C1200CBE141572DE">
Patrick Müller Amber Collection (Zweibrücken,
<collectingCountry id="7A39C68FD5EEC98B5ED4F8DB9175CD97" name="Germany">Germany</collectingCountry>
)
<collectingCountry id="2B8AEF989BDA7E7C209F433EEAAA7F9F" name="Mexico">MEX</collectingCountry>
70,
<collectingCountry id="0C4EA43523617ADF20E0463E447B7E7A" name="Mexico">MEX</collectingCountry>
71.
</paragraph>
</subSubSection>
<subSubSection id="SECID0EA3BG" type="etymology">
<paragraph id="E652F1DE98447756578D1D3D0FA96BD4">
<heading id="74F888F1CD95C0AA42704B85B0D68874" reason="title">Etymology.</heading>
</paragraph>
<paragraph id="1FBB07DEF077F1AFD24256397FE36506">The specific epithet honors the amber collector Patrick Müller who has generously supported our research by providing numerous amber fossils for study, including all known specimens of this species.</paragraph>
</subSubSection>
<subSubSection id="SECID0EF3BG" type="Age and stratigraphic level">
<paragraph id="3141EE138D74A0CFB17AF31C0F274791">
<heading id="F419859460171CF6BC7113467844D393" reason="title">Age and stratigraphic level.</heading>
</paragraph>
<paragraph id="83FCC6D9D857A69F5F247D96B67D6505">
1523 Ma, Langhian Aquitanian (early to middle Miocene), Simojovel,
<collectingRegion id="47350C7B0ED2FD35C55E09AA3FD11F7D" country="Mexico" name="Chiapas">Chiapas</collectingRegion>
,
<collectingCountry id="4722ED689BBB0C463E58E582A9774E34" name="Mexico">Mexico</collectingCountry>
.
</paragraph>
</subSubSection>
<subSubSection id="SECID0EU3BG" type="diagnosis">
<paragraph id="BB10F2B540A83740325BC5D0F91F69EA">
<heading id="D744DAA19F07D1F13BB18A71D0322886" reason="title">Diagnosis.</heading>
</paragraph>
<paragraph id="196FA3D54A836C15E2E99AEE3736F702">
Incubously foliated liverwort with a ventral merophyte at least five cells wide; flagelliform branches present on lower parts of the shoots; lateral leaves conduplicate-bilobed; dorsal lobe convex, asymmetrically oblong-ovate with a rounded to obtuse apex, median lobe cells mostly elongate with distinctly cordate trigones; ventral lobule
<taxonomicName id="3A9AC5268D509FDCA22485F30A0F96B0" class="Jungermanniopsida" family="Lejeuneaceae" genus="Lejeunea" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="genus">
<emphasis id="D3410F91027C33DC6E5BE4933C9BFBB7" italics="true">Lejeunea</emphasis>
</taxonomicName>
- type, ovate-rectangular with one hooked apical tooth and a truncate apex; underleaves suborbicular to rounded quadrate to obovate with a rounded to emarginate, recurved apex and entire, mostly revolute margins.
</paragraph>
</subSubSection>
<subSubSection id="SECID0ET4BG" type="description">
<paragraph id="3E9164706587067271E3524FF2C2C3A1">
<heading id="6C352C121C2A34E0436A00B903F6A795" reason="title">Description.</heading>
</paragraph>
<paragraph id="94EB701BC2774F19283D1E36B5BA2A67">
Gametophyte fragments yellowish to dark reddish brown,
<quantity id="C460B555FA9956FD0AEE7E6F4E49A9F2" metricMagnitude="-3" metricUnit="m" metricValue="3.87" metricValueMax="5.04" metricValueMin="2.7" unit="mm" value="3.87" valueMax="5.04" valueMin="2.7">2.75.04 mm</quantity>
long, main shoots
<quantity id="A438C6AA2B3455D0B3BF4DF5D06F283D" metricMagnitude="-4" metricUnit="m" metricValue="8.4" metricValueMax="11.6" metricValueMin="5.2" unit="mm" value="0.84" valueMax="1.16" valueMin="0.52">0.521.16 mm</quantity>
wide with leaves (Figs
<figureCitation id="FB31616CBB8FAB572C71D615792E02A3" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Thysananthus patrickmuelleri sp. nov. from Mexican amber, holotype (GZG. BST. 22086). A. Upper part of the gametophyte in dorsal view; B. Lower part of the gametophyte with a short Lejeunea - type flagelliform branch (indicated by white arrow) in dorsal view; C. Upper part of the gametophyte in ventral view; D. Flagelliform branch in ventral view; E. Underleaves and leaf lobules with single apical teeth; F. Lobe cells with cordate trigones." figureDoi="10.3897/fr.28.137758.figure4" httpUri="https://binary.pensoft.net/fig/1264816">4 A C</figureCitation>
,
<figureCitation id="4D65BD852A1045207457A892EBCADAF5" captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Thysananthus patrickmuelleri sp. nov. from Mexican amber (Patrick Müller amber collection, MEX 70). A. Gametophyte with erect-spreading leaves in dorsal view; B. Same specimen in ventral view; C. Upper part of the shoot in ventral view; D. Apical part of a ventral leaf lobule with a hooked tooth (indicated by white arrow); E. Lobe cells, mostly collapsed." figureDoi="10.3897/fr.28.137758.figure5" httpUri="https://binary.pensoft.net/fig/1264817">5 A C</figureCitation>
,
<figureCitation id="4C6C37862D64A5C82C68FF9AAF7DE9B6" captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. A E. Thysananthus patrickmuelleri sp. nov. (A E, Patrick Müller amber collection, MEX 71) and a representative of Ptychanthoideae (F, Stuttgart State Museum of Natural History, Mx 357) from Mexican amber. A. Branched gametophyte of Thysananthus patrickmuelleri in dorso-lateral view; B. Left shoot with erect-spreading leaves similar to MEX 70 and right shoot with spreading leaves similar to holotype GZG. BST. 22086; C. Lower part with several flagelliform branches, the one on the left becoming main shoot-like; D. Flagelliform branch in ventral view; E. Detail of the main shoot in ventral view; F. Ptychanthoideae sp., gametophyte in ventral view." figureDoi="10.3897/fr.28.137758.figure6" httpUri="https://binary.pensoft.net/fig/1264818">6 A E</figureCitation>
).
<emphasis id="F32BF9EB19DAC07E97482400AEED4297" bold="true" italics="true">Branching</emphasis>
irregular,
<taxonomicName id="1DEE1C4328A6400FEBA805429A4C76AA" class="Jungermanniopsida" family="Lejeuneaceae" genus="Lejeunea" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="genus">
<emphasis id="D0C91DB5964798D768B9833A2603C73A" italics="true">Lejeunea</emphasis>
</taxonomicName>
- type, primary branches either similar to main shoot, or flagelliform and microphyllous (Figs
<figureCitation id="01A6C4262D4E28386A333BF9AF32848A" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Thysananthus patrickmuelleri sp. nov. from Mexican amber, holotype (GZG. BST. 22086). A. Upper part of the gametophyte in dorsal view; B. Lower part of the gametophyte with a short Lejeunea - type flagelliform branch (indicated by white arrow) in dorsal view; C. Upper part of the gametophyte in ventral view; D. Flagelliform branch in ventral view; E. Underleaves and leaf lobules with single apical teeth; F. Lobe cells with cordate trigones." figureDoi="10.3897/fr.28.137758.figure4" httpUri="https://binary.pensoft.net/fig/1264816">4 B, D</figureCitation>
,
<figureCitation id="56A59795CD8E52D0B6F28F4292F6B459" captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. A E. Thysananthus patrickmuelleri sp. nov. (A E, Patrick Müller amber collection, MEX 71) and a representative of Ptychanthoideae (F, Stuttgart State Museum of Natural History, Mx 357) from Mexican amber. A. Branched gametophyte of Thysananthus patrickmuelleri in dorso-lateral view; B. Left shoot with erect-spreading leaves similar to MEX 70 and right shoot with spreading leaves similar to holotype GZG. BST. 22086; C. Lower part with several flagelliform branches, the one on the left becoming main shoot-like; D. Flagelliform branch in ventral view; E. Detail of the main shoot in ventral view; F. Ptychanthoideae sp., gametophyte in ventral view." figureDoi="10.3897/fr.28.137758.figure6" httpUri="https://binary.pensoft.net/fig/1264818">6 C, D</figureCitation>
); main shoot-like branches
<quantity id="16956DD90EF6CA2A3E5EE01CDC39031A" metricMagnitude="-3" metricUnit="m" metricValue="3.03" metricValueMax="4.8" metricValueMin="1.26" unit="mm" value="3.03" valueMax="4.8" valueMin="1.26">1.264.8 mm</quantity>
long,
<quantity id="6702535B5E33709B47EBBAD5C4CCD93F" metricMagnitude="-4" metricUnit="m" metricValue="9.049999999999999" metricValueMax="14.0" metricValueMin="4.1" unit="mm" value="0.9049999999999999" valueMax="1.4" valueMin="0.41">0.411.4 mm</quantity>
wide with leaves; flagelliform branches
<quantity id="3441142B0056D247B6FADD751D6F3AA9" metricMagnitude="-4" metricUnit="m" metricValue="2.2" metricValueMax="2.4" metricValueMin="2.0" unit="mm" value="0.22" valueMax="0.24" valueMin="0.2">0.20.24 mm</quantity>
wide; one secondary microphyllous branch becoming main shoot-like on upper part.
<emphasis id="9633E636E88B7D170DC0DF846EFF0EC7" bold="true" italics="true">Stem</emphasis>
dark reddish brown to blackish, 4070 µm in diameter; cortical cells possibly thick-walled and elongated [mostly obscured by underleaves], ventral merophyte ca. 5 cells wide; stem on large branches 4060 µm in diameter, on flagelliform branches ca. 30 µm in diameter.
<emphasis id="0D181B4FD481D0B3869B2F303D8D130D" bold="true" italics="true">Lateral leaves</emphasis>
incubous, densely imbricate, alternate, widely spreading to erect-spreading or erect-appressed (especially on upper shoot parts), conduplicate-bilobed with large dorsal lobe and a
<taxonomicName id="87AA535CDF1C0EC7F444CBE39D0F7E51" class="Jungermanniopsida" family="Lejeuneaceae" genus="Lejeunea" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="genus">
<emphasis id="9AB0E49831C204C35C626A188B69B117" italics="true">Lejeunea</emphasis>
</taxonomicName>
- type ventral lobule folded against the lobe (Figs
<figureCitation id="69A8979D17EE66A3ABE54BA332676F5E" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Thysananthus patrickmuelleri sp. nov. from Mexican amber, holotype (GZG. BST. 22086). A. Upper part of the gametophyte in dorsal view; B. Lower part of the gametophyte with a short Lejeunea - type flagelliform branch (indicated by white arrow) in dorsal view; C. Upper part of the gametophyte in ventral view; D. Flagelliform branch in ventral view; E. Underleaves and leaf lobules with single apical teeth; F. Lobe cells with cordate trigones." figureDoi="10.3897/fr.28.137758.figure4" httpUri="https://binary.pensoft.net/fig/1264816">4 C, E</figureCitation>
,
<figureCitation id="52B3689985AD0331AF4157F6CF49805F" captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Thysananthus patrickmuelleri sp. nov. from Mexican amber (Patrick Müller amber collection, MEX 70). A. Gametophyte with erect-spreading leaves in dorsal view; B. Same specimen in ventral view; C. Upper part of the shoot in ventral view; D. Apical part of a ventral leaf lobule with a hooked tooth (indicated by white arrow); E. Lobe cells, mostly collapsed." figureDoi="10.3897/fr.28.137758.figure5" httpUri="https://binary.pensoft.net/fig/1264817">5 C</figureCitation>
,
<figureCitation id="E4B27247FB7AF7BA724233EDEC4C594E" captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. A E. Thysananthus patrickmuelleri sp. nov. (A E, Patrick Müller amber collection, MEX 71) and a representative of Ptychanthoideae (F, Stuttgart State Museum of Natural History, Mx 357) from Mexican amber. A. Branched gametophyte of Thysananthus patrickmuelleri in dorso-lateral view; B. Left shoot with erect-spreading leaves similar to MEX 70 and right shoot with spreading leaves similar to holotype GZG. BST. 22086; C. Lower part with several flagelliform branches, the one on the left becoming main shoot-like; D. Flagelliform branch in ventral view; E. Detail of the main shoot in ventral view; F. Ptychanthoideae sp., gametophyte in ventral view." figureDoi="10.3897/fr.28.137758.figure6" httpUri="https://binary.pensoft.net/fig/1264818">6 E</figureCitation>
), insertion line J-shaped.
<emphasis id="B43B5D6251D5BB57E1AB34976ECC8A9B" bold="true" italics="true">Dorsal lobe</emphasis>
convex, asymmetrically oblong-ovate, on upper shoot parts falcate, 480770 µm long × 220300 µm wide in the middle, length: width ratio 1.43.5: 1, margin entire, postical margin nearly plane or slightly incurved for up to 0.7 × the lobe length, exterior to keel first weakly then abruptly arched towards apex, apex rounded to obtuse, antical margin regularly arched, lobes overlapping stem 0.52 × the stem width beyond the farther edge of the stem, dorsal base not visible; lobes on larger branches very similar to main shoot, short ovate to oblong-ovate, slightly falcate to straight, 230840 µm long × 160430 µm wide in the middle, length: width ratio 1.42.1: 1; lobes on flagelliform branches erect-spreading to erect, 150160 µm long × 70110 µm wide in the middle, length: width ratio 1.52.1: 1.
<emphasis id="8149B9CFB93C43B9E1FC86008C43D4DE" bold="true" italics="true">Lobe cells</emphasis>
rectangular to hexagonal (Figs
<figureCitation id="D487E1BCDE00C0918AF4EAE54212F919" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Thysananthus patrickmuelleri sp. nov. from Mexican amber, holotype (GZG. BST. 22086). A. Upper part of the gametophyte in dorsal view; B. Lower part of the gametophyte with a short Lejeunea - type flagelliform branch (indicated by white arrow) in dorsal view; C. Upper part of the gametophyte in ventral view; D. Flagelliform branch in ventral view; E. Underleaves and leaf lobules with single apical teeth; F. Lobe cells with cordate trigones." figureDoi="10.3897/fr.28.137758.figure4" httpUri="https://binary.pensoft.net/fig/1264816">4 F</figureCitation>
,
<figureCitation id="31B9AF3CA57DE2AE8F745EFAD8F0EE4B" captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Thysananthus patrickmuelleri sp. nov. from Mexican amber (Patrick Müller amber collection, MEX 70). A. Gametophyte with erect-spreading leaves in dorsal view; B. Same specimen in ventral view; C. Upper part of the shoot in ventral view; D. Apical part of a ventral leaf lobule with a hooked tooth (indicated by white arrow); E. Lobe cells, mostly collapsed." figureDoi="10.3897/fr.28.137758.figure5" httpUri="https://binary.pensoft.net/fig/1264817">5 D, E</figureCitation>
,
<figureCitation id="348D19B62E343AD18A164421508F29D3" captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. A E. Thysananthus patrickmuelleri sp. nov. (A E, Patrick Müller amber collection, MEX 71) and a representative of Ptychanthoideae (F, Stuttgart State Museum of Natural History, Mx 357) from Mexican amber. A. Branched gametophyte of Thysananthus patrickmuelleri in dorso-lateral view; B. Left shoot with erect-spreading leaves similar to MEX 70 and right shoot with spreading leaves similar to holotype GZG. BST. 22086; C. Lower part with several flagelliform branches, the one on the left becoming main shoot-like; D. Flagelliform branch in ventral view; E. Detail of the main shoot in ventral view; F. Ptychanthoideae sp., gametophyte in ventral view." figureDoi="10.3897/fr.28.137758.figure6" httpUri="https://binary.pensoft.net/fig/1264818">6 E</figureCitation>
), elongated at the lobe base and in the middle, slightly shorter to (sub) isodiametric near apex, up to 3 × as long as wide [often collapsed and deformed], basal cells 2035 µm long × 7.520 µm wide, median cells 1535 µm long × 7.517.5 µm wide, apical cells 7.520 µm long × 7.515 µm wide; cell walls thin, trigones cordate, not coalesced, few intermediate thickenings present.
<emphasis id="A47E74BAC11FB8B18A8F903F0CF26266" bold="true" italics="true">Ventral lobule</emphasis>
ovate-rectangular, lower part inflated, free antical margin not inflexed (Figs
<figureCitation id="03A8B1C0DDB4A49C395C648A2C155382" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Thysananthus patrickmuelleri sp. nov. from Mexican amber, holotype (GZG. BST. 22086). A. Upper part of the gametophyte in dorsal view; B. Lower part of the gametophyte with a short Lejeunea - type flagelliform branch (indicated by white arrow) in dorsal view; C. Upper part of the gametophyte in ventral view; D. Flagelliform branch in ventral view; E. Underleaves and leaf lobules with single apical teeth; F. Lobe cells with cordate trigones." figureDoi="10.3897/fr.28.137758.figure4" httpUri="https://binary.pensoft.net/fig/1264816">4 E</figureCitation>
,
<figureCitation id="4C26C42A2847D8736F2FDD9023162ECD" captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Thysananthus patrickmuelleri sp. nov. from Mexican amber (Patrick Müller amber collection, MEX 70). A. Gametophyte with erect-spreading leaves in dorsal view; B. Same specimen in ventral view; C. Upper part of the shoot in ventral view; D. Apical part of a ventral leaf lobule with a hooked tooth (indicated by white arrow); E. Lobe cells, mostly collapsed." figureDoi="10.3897/fr.28.137758.figure5" httpUri="https://binary.pensoft.net/fig/1264817">5 C, D</figureCitation>
,
<figureCitation id="D80B12F0D9575E3BA1C261E6EB4BEC25" captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. A E. Thysananthus patrickmuelleri sp. nov. (A E, Patrick Müller amber collection, MEX 71) and a representative of Ptychanthoideae (F, Stuttgart State Museum of Natural History, Mx 357) from Mexican amber. A. Branched gametophyte of Thysananthus patrickmuelleri in dorso-lateral view; B. Left shoot with erect-spreading leaves similar to MEX 70 and right shoot with spreading leaves similar to holotype GZG. BST. 22086; C. Lower part with several flagelliform branches, the one on the left becoming main shoot-like; D. Flagelliform branch in ventral view; E. Detail of the main shoot in ventral view; F. Ptychanthoideae sp., gametophyte in ventral view." figureDoi="10.3897/fr.28.137758.figure6" httpUri="https://binary.pensoft.net/fig/1264818">6 E</figureCitation>
), 0.30.4 × the length of the lobe, 200310 µm long × 80200 µm wide in the middle, length: width ratio 1.62.9: 1, free antical margin nearly straight to slightly arched, free lateral margin truncate and mostly forming an angle of ca. 90 ° with the postical lobe margin, apex with a single, often hooked tooth (Figs
<figureCitation id="0E5192619403290171853C15EF19EE71" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Thysananthus patrickmuelleri sp. nov. from Mexican amber, holotype (GZG. BST. 22086). A. Upper part of the gametophyte in dorsal view; B. Lower part of the gametophyte with a short Lejeunea - type flagelliform branch (indicated by white arrow) in dorsal view; C. Upper part of the gametophyte in ventral view; D. Flagelliform branch in ventral view; E. Underleaves and leaf lobules with single apical teeth; F. Lobe cells with cordate trigones." figureDoi="10.3897/fr.28.137758.figure4" httpUri="https://binary.pensoft.net/fig/1264816">4 E</figureCitation>
,
<figureCitation id="0DFF913433B44083B3B22117404346E3" captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Thysananthus patrickmuelleri sp. nov. from Mexican amber (Patrick Müller amber collection, MEX 70). A. Gametophyte with erect-spreading leaves in dorsal view; B. Same specimen in ventral view; C. Upper part of the shoot in ventral view; D. Apical part of a ventral leaf lobule with a hooked tooth (indicated by white arrow); E. Lobe cells, mostly collapsed." figureDoi="10.3897/fr.28.137758.figure5" httpUri="https://binary.pensoft.net/fig/1264817">5 D</figureCitation>
), 1015 µm (12 cells) long × 1015 µm (12 cells) wide at base, hyaline papilla not seen, keel curved, leaves slightly emarginate at end of keel; no appendages at the base of the lobule or the keel seen; lobules on larger branches 0.30.5 × as long as lobes, 110310 µm long × 70160 µm wide in the middle, length: width ratio 1.32.4: 1; lobules on flagelliform branches 0.50.7 × as long as lobe, 90110 µm long × 6080 µm wide in the middle, length: width ratio 1.31.8: 1.
<emphasis id="3CAE54B326896FE8A9835F5513E2E675" bold="true" italics="true">Underleaves</emphasis>
imbricate to contiguous, symmetrical, suborbicular to rounded quadrate to obovate, generally wider than long to as long as wide (Figs
<figureCitation id="9C852EABE82EBB61CB41CEA3A7434520" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Thysananthus patrickmuelleri sp. nov. from Mexican amber, holotype (GZG. BST. 22086). A. Upper part of the gametophyte in dorsal view; B. Lower part of the gametophyte with a short Lejeunea - type flagelliform branch (indicated by white arrow) in dorsal view; C. Upper part of the gametophyte in ventral view; D. Flagelliform branch in ventral view; E. Underleaves and leaf lobules with single apical teeth; F. Lobe cells with cordate trigones." figureDoi="10.3897/fr.28.137758.figure4" httpUri="https://binary.pensoft.net/fig/1264816">4 C, E</figureCitation>
,
<figureCitation id="8A7BA77DCC8B1697282981A2CF760A5C" captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Thysananthus patrickmuelleri sp. nov. from Mexican amber (Patrick Müller amber collection, MEX 70). A. Gametophyte with erect-spreading leaves in dorsal view; B. Same specimen in ventral view; C. Upper part of the shoot in ventral view; D. Apical part of a ventral leaf lobule with a hooked tooth (indicated by white arrow); E. Lobe cells, mostly collapsed." figureDoi="10.3897/fr.28.137758.figure5" httpUri="https://binary.pensoft.net/fig/1264817">5 C</figureCitation>
,
<figureCitation id="918A3D1E8B32249BBAE3232189F7B56E" captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. A E. Thysananthus patrickmuelleri sp. nov. (A E, Patrick Müller amber collection, MEX 71) and a representative of Ptychanthoideae (F, Stuttgart State Museum of Natural History, Mx 357) from Mexican amber. A. Branched gametophyte of Thysananthus patrickmuelleri in dorso-lateral view; B. Left shoot with erect-spreading leaves similar to MEX 70 and right shoot with spreading leaves similar to holotype GZG. BST. 22086; C. Lower part with several flagelliform branches, the one on the left becoming main shoot-like; D. Flagelliform branch in ventral view; E. Detail of the main shoot in ventral view; F. Ptychanthoideae sp., gametophyte in ventral view." figureDoi="10.3897/fr.28.137758.figure6" httpUri="https://binary.pensoft.net/fig/1264818">6 E</figureCitation>
), 36 × wider than the stem, 150310 µm long × 160390 µm wide, length: width ratio 0.61: 1; slightly squarrose, concave with recurved margins to occasionally nearly plane, apex recurved, rounded to truncate to emarginate, margins entire, insertion line arched, base auriculate, not adnate to lateral leaf; underleaves on larger branches more often nearly plane, on lower parts slightly longer than wide, 160280 µm long × 130360 µm wide, length: width ratio 0.71.2: 1, apex rounded, often not recurved; underleaves on flagelliform branches not clearly visible, possibly ovate.
<emphasis id="64B776E65F95CFDB9196D2BA72FC3B7D" bold="true" italics="true">Rhizoids</emphasis>
in bundles near underleaf insertion, bundles up to 230 µm long, rhizoids ca. 10 µm in diameter with truncate and ampliate tips.
<emphasis id="702E3DF76FE9E5FC583BBF1F2CB600B1" bold="true">
<emphasis id="3F63AD9996B31CBBF1C285AA723A580E" bold="true" italics="true">Sterile</emphasis>
.
</emphasis>
</paragraph>
<caption id="90E671A5CB455F4DCC2EBA47DC87D298" ID-DOI="10.3897/fr.28.137758.figure4" ID-arpha="1A852CFB-09E3-5848-9DF5-AA23E9BF78B0" httpUri="https://binary.pensoft.net/fig/1264816" startId="F4">
<paragraph id="DEB333888026761B3313745ACC09795F">
<label id="686C147D80E70CAD1514D810FA2FF92A">Figure 4.</label>
</paragraph>
<paragraph id="60C4CF062D84B994E9A6C17FFA8C2C0C">
<taxonomicName id="CDCA92A9403A8F964F3F6686312592C6" authorityName="K. Feldberg, Gradst., Schaf. - Verw., Mamontov" authorityYear="2025" class="Jungermanniopsida" family="Lejeuneaceae" genus="Thysananthus" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="patrickmuelleri" status="sp. nov.">
<emphasis id="1FA3FFC1E63ACC5A93203D5015913C90" italics="true">Thysananthus patrickmuelleri</emphasis>
</taxonomicName>
<taxonomicNameLabel id="BCD73C503E4FE5FC76D2DA23C3A6E320" rank="species">sp. nov.</taxonomicNameLabel>
from Mexican amber, holotype (
<specimenCode id="74A474F398F58772B293B64D864A9920">
<abbrev content-type="institution" id="ABBRID0EZEAI" xlink_title="Geoscientific Collection of the University of Göttingen">GZG</abbrev>
.BST.22086
</specimenCode>
).
<emphasis id="D0EC78FEEC355FF74F1E0AA050CB422B" bold="true">A.</emphasis>
Upper part of the gametophyte in dorsal view;
<emphasis id="C97A6CEA57ADE308FB439B928D62A8F3" bold="true">B.</emphasis>
Lower part of the gametophyte with a short
<taxonomicName id="63830A1C7B96B884ED0FE5AE22BC79D4" class="Jungermanniopsida" family="Lejeuneaceae" genus="Lejeunea" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="genus">
<emphasis id="072E77D680D28E2CEE79C989E88F7FB6" italics="true">Lejeunea</emphasis>
</taxonomicName>
- type flagelliform branch (indicated by white arrow) in dorsal view;
<emphasis id="F6DA68DA4669DBF14459C50543E3BA95" bold="true">C.</emphasis>
Upper part of the gametophyte in ventral view;
<emphasis id="F800B74DEE88F56293E5E6339F42CAF9" bold="true">D.</emphasis>
Flagelliform branch in ventral view;
<emphasis id="D493FC76D9C67FB8A510BF85461D3341" bold="true">E.</emphasis>
Underleaves and leaf lobules with single apical teeth;
<emphasis id="4781FAEAA60254FA46AF07AFF3CE56B3" bold="true">F.</emphasis>
Lobe cells with cordate trigones.
</paragraph>
</caption>
<caption id="705C666125208637446CA1FF04E894E6" ID-DOI="10.3897/fr.28.137758.figure5" ID-arpha="64066B06-1EEB-5E0A-9D3E-FD0C01EAFB48" httpUri="https://binary.pensoft.net/fig/1264817" startId="F5">
<paragraph id="916E083CD6EB5C73A7FB2BC75310D752">
<label id="ED987725FCDCF0CBE322C157A5BFCDB2">Figure 5.</label>
</paragraph>
<paragraph id="CC0A14F98483373D179FDFAA27760CE5">
<taxonomicName id="0AAB81763B62F19C82F914DD9D7F2F3D" authorityName="K. Feldberg, Gradst., Schaf. - Verw., Mamontov" authorityYear="2025" class="Jungermanniopsida" family="Lejeuneaceae" genus="Thysananthus" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="patrickmuelleri" status="sp. nov.">
<emphasis id="C220EA150453CF3C063B70368D4C7803" italics="true">Thysananthus patrickmuelleri</emphasis>
</taxonomicName>
<taxonomicNameLabel id="590C6D2DBD5852183612F7DBE4B25A03" rank="species">sp. nov.</taxonomicNameLabel>
from Mexican amber (Patrick Müller amber collection, MEX 70).
<emphasis id="1CE975E1882E54B5B6CD2DEB0217B9B0" bold="true">A.</emphasis>
Gametophyte with erect-spreading leaves in dorsal view;
<emphasis id="773481DF86E4E66C460639D5EF92B753" bold="true">B.</emphasis>
Same specimen in ventral view;
<emphasis id="696E46F9203267BB96F9AC0BF50DBDE4" bold="true">C.</emphasis>
Upper part of the shoot in ventral view;
<emphasis id="9EF250FE097418762C7E91F175F21B09" bold="true">D.</emphasis>
Apical part of a ventral leaf lobule with a hooked tooth (indicated by white arrow);
<emphasis id="E6F8C7BDC4849CDC851D3AD726EDFC91" bold="true">E.</emphasis>
Lobe cells, mostly collapsed.
</paragraph>
</caption>
<caption id="04D4E86F4075FCEB7BA51807B5F02CA2" ID-DOI="10.3897/fr.28.137758.figure6" ID-arpha="93FCB0E8-DE53-50E3-820E-50FDE95D8CAD" httpUri="https://binary.pensoft.net/fig/1264818" startId="F6">
<paragraph id="68D1CB9547C18615EB8ECBCA4C0148F6">
<label id="E59406C5D8852221B0442091425C48E8">Figure 6.</label>
</paragraph>
<paragraph id="038D3E9021D0AAF6527569273A839BED">
<emphasis id="26BADFFCAC2A718EB2FC303514B0841B" bold="true">A E.</emphasis>
<taxonomicName id="9698031A6E9C76FB799763E9997EE93F" authorityName="K. Feldberg, Gradst., Schaf. - Verw., Mamontov" authorityYear="2025" class="Jungermanniopsida" family="Lejeuneaceae" genus="Thysananthus" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="patrickmuelleri" status="sp. nov.">
<emphasis id="5469A0FEC1CBE8836EFADF8776089313" italics="true">Thysananthus patrickmuelleri</emphasis>
</taxonomicName>
<taxonomicNameLabel id="CDAF5BEB01C6746CD853C64A1AA2666E" rank="species">sp. nov.</taxonomicNameLabel>
(
<emphasis id="30F209E3B8B3369D55A4AEE78E87E720" bold="true">A E</emphasis>
, Patrick Müller amber collection, MEX 71) and a representative of
<taxonomicName id="B725FACC48C889EB7AA28A8B43D18132" class="Jungermanniopsida" family="Lejeuneaceae" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="subFamily" subFamily="Ptychanthoideae">Ptychanthoideae</taxonomicName>
(
<emphasis id="F8E64A651411C6D50000D88B4BA1FB3B" bold="true">F</emphasis>
, Stuttgart State Museum of Natural History, Mx 357) from Mexican amber.
<emphasis id="2E4AB8CA280051835B3D4BC08008FDD7" bold="true">A.</emphasis>
Branched gametophyte of
<taxonomicName id="6F6AE766339A16F4DA7D610D260FCD3A" authorityName="K. Feldberg, Gradst., Schaf. - Verw., Mamontov" authorityYear="2025" class="Jungermanniopsida" family="Lejeuneaceae" genus="Thysananthus" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="patrickmuelleri">
<emphasis id="4CBFC1A959D9469B0FEB8D34B2AA1223" italics="true">Thysananthus patrickmuelleri</emphasis>
</taxonomicName>
in dorso-lateral view;
<emphasis id="CD8CBCD79015CE539C9D8C85D17B0A64" bold="true">B.</emphasis>
Left shoot with erect-spreading leaves similar to MEX 70 and right shoot with spreading leaves similar to holotype
<specimenCode id="D3B2F6258987F1749EF4B745C06218BA">
<abbrev content-type="institution" id="ABBRID0EOJAI" xlink_title="Geoscientific Collection of the University of Göttingen">GZG</abbrev>
.BST.22086
</specimenCode>
;
<emphasis id="824C75703CD84331DDBCC00957A1C3B9" bold="true">C.</emphasis>
Lower part with several flagelliform branches, the one on the left becoming main shoot-like;
<emphasis id="F7A0E60D9636411EBA011CBE2410A57E" bold="true">D.</emphasis>
Flagelliform branch in ventral view;
<emphasis id="D2F8E5576B88FCFAEB490E4ADED2D924" bold="true">E.</emphasis>
Detail of the main shoot in ventral view;
<emphasis id="27D5573A4DE807CC50FA986CB4E27CD3" bold="true">F.</emphasis>
<taxonomicName id="3FFDB1ACD688978C7D5E4E8AF3DF8DD2" class="Jungermanniopsida" family="Lejeuneaceae" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="subFamily" subFamily="Ptychanthoideae">Ptychanthoideae</taxonomicName>
sp., gametophyte in ventral view.
</paragraph>
</caption>
</subSubSection>
<subSubSection id="SECID0EPKAI" type="discussion">
<paragraph id="C0F4F3F813DA147FFB755F471D53F49E">
<heading id="A4F21F23DBCDF5EDCC737D6F5629DA7E" reason="title">Discussion.</heading>
</paragraph>
<paragraph id="C3EEF491BDB2DFCC21EDCE9B78652F25">
The conduplicate-bilobed leaves with
<taxonomicName id="82C69DA8A1E28D26FD481197215C57EA" class="Jungermanniopsida" family="Lejeuneaceae" genus="Lejeunea" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="genus">
<emphasis id="223B73129F9E85EA0507ABBEE998CDB1" italics="true">Lejeunea</emphasis>
</taxonomicName>
- type lobules and the presence of underleaves clearly identify this specimen as a member of
<taxonomicName id="FE4C3A4A27C5E806FB9E7EBE141AD81A" authorityName="Cavers" authorityYear="1910" class="Jungermanniopsida" family="Lejeuneaceae" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="family">Lejeuneaceae</taxonomicName>
(Figs
<figureCitation id="095DD8959B08C6864B735E86747531CC" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Thysananthus patrickmuelleri sp. nov. from Mexican amber, holotype (GZG. BST. 22086). A. Upper part of the gametophyte in dorsal view; B. Lower part of the gametophyte with a short Lejeunea - type flagelliform branch (indicated by white arrow) in dorsal view; C. Upper part of the gametophyte in ventral view; D. Flagelliform branch in ventral view; E. Underleaves and leaf lobules with single apical teeth; F. Lobe cells with cordate trigones." figureDoi="10.3897/fr.28.137758.figure4" httpUri="https://binary.pensoft.net/fig/1264816">4 C, E</figureCitation>
,
<figureCitation id="0F7930DE0587CEC28854FEDE7465D85C" captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Thysananthus patrickmuelleri sp. nov. from Mexican amber (Patrick Müller amber collection, MEX 70). A. Gametophyte with erect-spreading leaves in dorsal view; B. Same specimen in ventral view; C. Upper part of the shoot in ventral view; D. Apical part of a ventral leaf lobule with a hooked tooth (indicated by white arrow); E. Lobe cells, mostly collapsed." figureDoi="10.3897/fr.28.137758.figure5" httpUri="https://binary.pensoft.net/fig/1264817">5 A C</figureCitation>
,
<figureCitation id="C5CD39825296CBB34858A7F783866273" captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. A E. Thysananthus patrickmuelleri sp. nov. (A E, Patrick Müller amber collection, MEX 71) and a representative of Ptychanthoideae (F, Stuttgart State Museum of Natural History, Mx 357) from Mexican amber. A. Branched gametophyte of Thysananthus patrickmuelleri in dorso-lateral view; B. Left shoot with erect-spreading leaves similar to MEX 70 and right shoot with spreading leaves similar to holotype GZG. BST. 22086; C. Lower part with several flagelliform branches, the one on the left becoming main shoot-like; D. Flagelliform branch in ventral view; E. Detail of the main shoot in ventral view; F. Ptychanthoideae sp., gametophyte in ventral view." figureDoi="10.3897/fr.28.137758.figure6" httpUri="https://binary.pensoft.net/fig/1264818">6 E</figureCitation>
). The robust size, the undivided underleaves, and at least five cells wide ventral merophyte allow an assignment to subfamily
<taxonomicName id="5EE41CF941DE6A697F8363B886B5F94A" class="Jungermanniopsida" family="Lejeuneaceae" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="subFamily" subFamily="Ptychanthoideae">Ptychanthoideae</taxonomicName>
(Figs
<figureCitation id="FF5813B561992ED1D7504E1DEE9C0722" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Thysananthus patrickmuelleri sp. nov. from Mexican amber, holotype (GZG. BST. 22086). A. Upper part of the gametophyte in dorsal view; B. Lower part of the gametophyte with a short Lejeunea - type flagelliform branch (indicated by white arrow) in dorsal view; C. Upper part of the gametophyte in ventral view; D. Flagelliform branch in ventral view; E. Underleaves and leaf lobules with single apical teeth; F. Lobe cells with cordate trigones." figureDoi="10.3897/fr.28.137758.figure4" httpUri="https://binary.pensoft.net/fig/1264816">4 A, C</figureCitation>
,
<figureCitation id="A12370118C34A00E46DABB7196D0708D" captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Thysananthus patrickmuelleri sp. nov. from Mexican amber (Patrick Müller amber collection, MEX 70). A. Gametophyte with erect-spreading leaves in dorsal view; B. Same specimen in ventral view; C. Upper part of the shoot in ventral view; D. Apical part of a ventral leaf lobule with a hooked tooth (indicated by white arrow); E. Lobe cells, mostly collapsed." figureDoi="10.3897/fr.28.137758.figure5" httpUri="https://binary.pensoft.net/fig/1264817">5 A C</figureCitation>
,
<figureCitation id="7E87181781BFD7448107438B5EEFCBF5" captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. A E. Thysananthus patrickmuelleri sp. nov. (A E, Patrick Müller amber collection, MEX 71) and a representative of Ptychanthoideae (F, Stuttgart State Museum of Natural History, Mx 357) from Mexican amber. A. Branched gametophyte of Thysananthus patrickmuelleri in dorso-lateral view; B. Left shoot with erect-spreading leaves similar to MEX 70 and right shoot with spreading leaves similar to holotype GZG. BST. 22086; C. Lower part with several flagelliform branches, the one on the left becoming main shoot-like; D. Flagelliform branch in ventral view; E. Detail of the main shoot in ventral view; F. Ptychanthoideae sp., gametophyte in ventral view." figureDoi="10.3897/fr.28.137758.figure6" httpUri="https://binary.pensoft.net/fig/1264818">6 A, B, E</figureCitation>
). Although all specimens are sterile, the ovate, apically rounded dorsal lobes, the elongate median lobe cells with distinct cordate trigones, as well as the entire-margined underleaves, the probably thick-walled stem cells, and the presence of flagelliform branches on lower shoot parts are indicative of the genus
<taxonomicName id="25E8978F59829C8563D085523117B46F" class="Jungermanniopsida" family="Lejeuneaceae" genus="Thysananthus" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="genus">
<emphasis id="5D407330154775E9762B84D2643DA875" italics="true">Thysananthus</emphasis>
</taxonomicName>
(Figs
<figureCitation id="E9291EB67C0E9BE687CC3B1F62E9E728" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Thysananthus patrickmuelleri sp. nov. from Mexican amber, holotype (GZG. BST. 22086). A. Upper part of the gametophyte in dorsal view; B. Lower part of the gametophyte with a short Lejeunea - type flagelliform branch (indicated by white arrow) in dorsal view; C. Upper part of the gametophyte in ventral view; D. Flagelliform branch in ventral view; E. Underleaves and leaf lobules with single apical teeth; F. Lobe cells with cordate trigones." figureDoi="10.3897/fr.28.137758.figure4" httpUri="https://binary.pensoft.net/fig/1264816">4</figureCitation>
,
<figureCitation id="20C46BD35BE61478EBE484D346292403" captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Thysananthus patrickmuelleri sp. nov. from Mexican amber (Patrick Müller amber collection, MEX 70). A. Gametophyte with erect-spreading leaves in dorsal view; B. Same specimen in ventral view; C. Upper part of the shoot in ventral view; D. Apical part of a ventral leaf lobule with a hooked tooth (indicated by white arrow); E. Lobe cells, mostly collapsed." figureDoi="10.3897/fr.28.137758.figure5" httpUri="https://binary.pensoft.net/fig/1264817">5</figureCitation>
,
<figureCitation id="40CAB8BB10AFC4546707C5602E126E65" captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. A E. Thysananthus patrickmuelleri sp. nov. (A E, Patrick Müller amber collection, MEX 71) and a representative of Ptychanthoideae (F, Stuttgart State Museum of Natural History, Mx 357) from Mexican amber. A. Branched gametophyte of Thysananthus patrickmuelleri in dorso-lateral view; B. Left shoot with erect-spreading leaves similar to MEX 70 and right shoot with spreading leaves similar to holotype GZG. BST. 22086; C. Lower part with several flagelliform branches, the one on the left becoming main shoot-like; D. Flagelliform branch in ventral view; E. Detail of the main shoot in ventral view; F. Ptychanthoideae sp., gametophyte in ventral view." figureDoi="10.3897/fr.28.137758.figure6" httpUri="https://binary.pensoft.net/fig/1264818">6 A E</figureCitation>
).
</paragraph>
<paragraph id="140A7B307BA5680F0AA4E0D9CFC6CB04">
<taxonomicName id="EEDDF053145DE5D10ABD486A5632DD68" class="Jungermanniopsida" family="Lejeuneaceae" genus="Thysananthus" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="genus">
<emphasis id="7F796379D6640FE0BF7DF64A4FEC6BE4" italics="true">Thysananthus</emphasis>
</taxonomicName>
is a large genus with 30 extant species and several fossil taxa (
<bibRefCitation id="FED3B1AC8AFDE17231D537D01B70F433" author="Gradstein" firstAuthor="Gradstein" journalOrPublisher="Nova Hedwigia" pagination="353-374" refId="B31" refString="Gradstein SR (1993) New fossil Hepaticae preserved in amber of the Dominican Republic. Nova Hedwigia 57: 353374." title="New fossil Hepaticae preserved in amber of the Dominican Republic." volume="57" year="1993">Gradstein 1993</bibRefCitation>
;
<bibRefCitation id="444DD73676DDB372C26DFB99913CBB27" DOI="10.1127/0029-5035/2014/0206" author="Sukkharak and Gradstein" firstAuthor="Sukkharak" journalOrPublisher="Nova Hedwigia" pagination="279-345" refId="B130" refString="Sukkharak P, Gradstein SR (2014) A taxonomic revision of the genus Mastigolejeunea (Marchantiophyta: Lejeuneaceae). Nova Hedwigia 99: 279345. https://doi.org/10.1127/0029-5035/2014/0206" title="A taxonomic revision of the genus Mastigolejeunea (Marchantiophyta: Lejeuneaceae)." volume="99" year="2014">Sukkharak and Gradstein 2014</bibRefCitation>
,
<bibRefCitation id="4CEBD2A7064470A20FAC1BEC2D6A283B" DOI="10.11646/phytotaxa.326.2.1" author="Sukkharak and Gradstein" firstAuthor="Sukkharak" refId="B131" refString="Sukkharak P, Gradstein SR (2017) Phylogenetic study of Mastigolejeunea (Marchantiophyta: Lejeuneaceae) and an amended circumscription of the genus Thysananthus. Phytotaxa 326: 91107. https://doi.org/10.11646/phytotaxa.326.2.1" year="2017">2017</bibRefCitation>
;
<bibRefCitation id="E571297F84F6124FF1B5304AEC34D7AC" DOI="10.11646/phytotaxa.193.1.1" author="Sukkharak" firstAuthor="Sukkharak" journalOrPublisher="Phytotaxa" pagination="1-81" refId="B129" refString="Sukkharak P (2015) A systematic monograph of the genus Thysananthus (Lejeuneaceae, Marchantiophyta). Phytotaxa 193: 181. https://doi.org/10.11646/phytotaxa.193.1.1" title="A systematic monograph of the genus Thysananthus (Lejeuneaceae, Marchantiophyta)." volume="193" year="2015">Sukkharak 2015</bibRefCitation>
;
<bibRefCitation id="0280D7E83FEC03CFADD087CD6F670451" author="Wang" etAl="et al." firstAuthor="Wang" journalOrPublisher="Bryophytorum Bibliotheca" pagination="1-141" refId="B141" refString="Wang J, Zhu R-L, Gradstein SR (2016) Taxonomic revision of Lejeuneaceae subfamily Ptychanthoideae (Marchantiophyta) in China. Bryophytorum Bibliotheca 65: 1141. https://www.schweizerbart.de/publications/detail/isbn/9783443620370" title="Taxonomic revision of Lejeuneaceae subfamily Ptychanthoideae (Marchantiophyta) in China." url="https://www.schweizerbart.de/publications/detail/isbn/9783443620370" volume="65" year="2016">Wang et al. 2016</bibRefCitation>
;
<bibRefCitation id="9EE3778F397FDDFBD7FF0EA68BD5C51C" DOI="10.11646/bde.43.1.6" author="Feldberg K &amp; Gradstein SR &amp; Gröhn C &amp; Heinrichs J &amp; von Konrat M &amp; Mamontov YS &amp; Renner MAM &amp; Roth M &amp; Schäfer-Verwimp A &amp; Sukkharak P &amp; Schmidt AR" journalOrPublisher="Bryophyte Diversity and Evolution" pagination="14-71" refId="B23" refString="Feldberg K, Gradstein SR, Gröhn C, Heinrichs J, von Konrat M, Mamontov YS, Renner MAM, Roth M, Schäfer-Verwimp A, Sukkharak P, Schmidt AR (2021 a) Checklist of fossil liverworts suitable for calibrating phylogenetic reconstructions. Bryophyte Diversity and Evolution 43: 1471. https://doi.org/10.11646/bde.43.1.6" title="Checklist of fossil liverworts suitable for calibrating phylogenetic reconstructions." volume="43">Feldberg et al. 2021 a</bibRefCitation>
). It includes the former genus
<taxonomicName id="B7BEF21C31986F8DA0953A96DB76CEB1" class="Jungermanniopsida" family="Lejeuneaceae" genus="Mastigolejeunea" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="genus">
<emphasis id="0032FC0F38A60EE4E70AF3831EB07306" italics="true">Mastigolejeunea</emphasis>
</taxonomicName>
which was ranked at subgenus level based on molecular phylogenetic analyses and morphology (
<bibRefCitation id="AEA633586BF50CAA77D530EACCEA4EF4" DOI="10.11646/phytotaxa.326.2.1" author="Sukkharak and Gradstein" firstAuthor="Sukkharak" refId="B131" refString="Sukkharak P, Gradstein SR (2017) Phylogenetic study of Mastigolejeunea (Marchantiophyta: Lejeuneaceae) and an amended circumscription of the genus Thysananthus. Phytotaxa 326: 91107. https://doi.org/10.11646/phytotaxa.326.2.1" year="2017">Sukkharak and Gradstein 2017</bibRefCitation>
). The oldest fossil of the genus is
<taxonomicName id="8453F55449C1353B3FC02B9552C1482D" authority="(Gopp. &amp; Berendt) Sukkharak &amp; Gradst." authorityName="Sukkharak &amp; Gradst." baseAuthorityName="Gopp. &amp; Berendt" class="Jungermanniopsida" family="Lejeuneaceae" genus="Thysananthus" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="contortus">
<emphasis id="9016D2EE185D791382956F8F6886F173" italics="true">T. contortus</emphasis>
(Göpp. &amp; Berendt) Sukkharak &amp; Gradst.
</taxonomicName>
from Paleogene Baltic and Bitterfeld amber. This species has predominantly
<taxonomicName id="AF0FC61997CD05A2F4C47A28935C4E24" class="Jungermanniopsida" family="Frullaniaceae" genus="Frullania" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="genus">
<emphasis id="B4DA112267AC17986917C098BB648ED4" italics="true">Frullania</emphasis>
</taxonomicName>
- type branches, which become as vigorous as the main axis, and rarely
<taxonomicName id="E161271A0AD3E202766644777A592536" class="Jungermanniopsida" family="Lejeuneaceae" genus="Lejeunea" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="genus">
<emphasis id="E72F99D2440A82FFD4446FECAEBA0054" italics="true">Lejeunea</emphasis>
</taxonomicName>
- type branches, as well as four lobule teeth (
<bibRefCitation id="1A27E82C528F70C2AA6676E894AC3E2D" author="Grolle and Meister" firstAuthor="Grolle" journalOrPublisher="Weissdorn Verlag, Jena" refId="B50" refString="Grolle R, Meister K (2004) The Liverworts in Baltic and Bitterfeld Amber. Weissdorn Verlag, Jena, 91 pp." title="The Liverworts in Baltic and Bitterfeld Amber." year="2004">Grolle and Meister 2004</bibRefCitation>
, plates 14, 15, as
<taxonomicName id="8BC91AEE1671C74E7C4FEBFD994D0713" authority="(Gopp. &amp; Berendt) Gradst. &amp; Grolle" authorityName="Gradst. &amp; Grolle" baseAuthorityName="Gopp. &amp; Berendt" class="Jungermanniopsida" family="Lejeuneaceae" genus="Mastigolejeunea" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="contorta">
<emphasis id="64E64E1E25D1C19F52F5184676190AD5" italics="true">Mastigolejeunea contorta</emphasis>
(Göpp. &amp; Berendt) Gradst. &amp; Grolle
</taxonomicName>
;
<bibRefCitation id="85D8A0E81BB76AC2C8C2D95C0242E90B" DOI="10.11646/phytotaxa.326.2.1" author="Sukkharak and Gradstein" firstAuthor="Sukkharak" refId="B131" refString="Sukkharak P, Gradstein SR (2017) Phylogenetic study of Mastigolejeunea (Marchantiophyta: Lejeuneaceae) and an amended circumscription of the genus Thysananthus. Phytotaxa 326: 91107. https://doi.org/10.11646/phytotaxa.326.2.1" year="2017">Sukkharak and Gradstein 2017</bibRefCitation>
). More fossil species of
<taxonomicName id="B64A513E442C80A6AABFE4FA3F9397B8" class="Jungermanniopsida" family="Lejeuneaceae" genus="Thysananthus" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="genus">
<emphasis id="D38C4DD2C784DF176E8742DFA9BD7D43" italics="true">Thysananthus</emphasis>
</taxonomicName>
have been found in tropical Miocene ambers. The genus is known from Dominican amber with the extant species
<taxonomicName id="D84B2CC0E2E683F0FB1BC29B199B4562" authority="(Wilson &amp; Hook.) Sukkharak &amp; Gradst." authorityName="Sukkharak &amp; Gradst." baseAuthorityName="Wilson &amp; Hook." class="Jungermanniopsida" family="Lejeuneaceae" genus="Thysananthus" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="auriculatus">
<emphasis id="ECF71FD067B0721E6697DADD1976F91A" italics="true">Thysananthus auriculatus</emphasis>
(Wilson &amp; Hook.) Sukkharak &amp; Gradst.
</taxonomicName>
(
<bibRefCitation id="C9FB0F6F0BD9AAAAF06568EBA422E252" author="Gradstein" firstAuthor="Gradstein" journalOrPublisher="Nova Hedwigia" pagination="353-374" refId="B31" refString="Gradstein SR (1993) New fossil Hepaticae preserved in amber of the Dominican Republic. Nova Hedwigia 57: 353374." title="New fossil Hepaticae preserved in amber of the Dominican Republic." volume="57" year="1993">Gradstein 1993</bibRefCitation>
, as
<taxonomicName id="A4363A817C9037E138735BA962C54D65" authority="(Wilson &amp; Hook.) Steph." authorityName="Steph." baseAuthorityName="Wilson &amp; Hook." class="Jungermanniopsida" family="Lejeuneaceae" genus="Mastigolejeunea" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="auriculata">
<emphasis id="40A7AFAE2CA8D247ABCB014D80FAB877" italics="true">Mastigolejeunea auriculata</emphasis>
(Wilson &amp; Hook.) Steph.
</taxonomicName>
;
<bibRefCitation id="74DF4D4FBD334E9B21EDBF3CB72378F5" DOI="10.11646/phytotaxa.326.2.1" author="Sukkharak and Gradstein" firstAuthor="Sukkharak" refId="B131" refString="Sukkharak P, Gradstein SR (2017) Phylogenetic study of Mastigolejeunea (Marchantiophyta: Lejeuneaceae) and an amended circumscription of the genus Thysananthus. Phytotaxa 326: 91107. https://doi.org/10.11646/phytotaxa.326.2.1" year="2017">Sukkharak and Gradstein 2017</bibRefCitation>
) as well as the extinct species
<taxonomicName id="8EC99290A2EFF0A0D778F50B616CD9EB" authority="(Gradst.) Sukkharak &amp; Gradst." authorityName="Sukkharak &amp; Gradst." baseAuthorityName="Gradst." class="Jungermanniopsida" family="Lejeuneaceae" genus="Thysananthus" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="bidentulus">
<emphasis id="86D20DD63BA45DB344AE6BCBB72F3114" italics="true">T. bidentulus</emphasis>
(Gradst.) Sukkharak &amp; Gradst.
</taxonomicName>
(
<bibRefCitation id="9F6EB0E4C6F83D31B5CF9AB92A15A656" author="Gradstein" firstAuthor="Gradstein" journalOrPublisher="Nova Hedwigia" pagination="353-374" refId="B31" refString="Gradstein SR (1993) New fossil Hepaticae preserved in amber of the Dominican Republic. Nova Hedwigia 57: 353374." title="New fossil Hepaticae preserved in amber of the Dominican Republic." volume="57" year="1993">Gradstein 1993</bibRefCitation>
, as
<taxonomicName id="656F656F49D21B95F82C15B71C4B3DB3" authority="Gradst." authorityName="Gradst." class="Jungermanniopsida" family="Lejeuneaceae" genus="Mastigolejeunea" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="bidentula">
<emphasis id="00D7FBB1C3FDDF94F38C6BE5E1EB4B94" italics="true">Mastigolejeunea bidentula</emphasis>
Gradst.
</taxonomicName>
;
<bibRefCitation id="4BBDF02C066E7D6EE13085D07B8B8E93" DOI="10.11646/phytotaxa.326.2.1" author="Sukkharak and Gradstein" firstAuthor="Sukkharak" refId="B131" refString="Sukkharak P, Gradstein SR (2017) Phylogenetic study of Mastigolejeunea (Marchantiophyta: Lejeuneaceae) and an amended circumscription of the genus Thysananthus. Phytotaxa 326: 91107. https://doi.org/10.11646/phytotaxa.326.2.1" year="2017">Sukkharak and Gradstein 2017</bibRefCitation>
) and
<taxonomicName id="C213F8DC112B3536F991242D06DE2818" authority="N. - N. Yu &amp; Gradst." authorityName="N. - N. Yu &amp; Gradst." class="Jungermanniopsida" family="Lejeuneaceae" genus="Thysananthus" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="weiweianus">
<emphasis id="C2C7FA1FE3EA5BD14A47D061E681F955" italics="true">T. weiweianus</emphasis>
N. - N. Yu &amp; Gradst.
</taxonomicName>
(
<bibRefCitation id="6ECC02D05DEC87233E64DE3E929CE644" author="Yu" etAl="et al." firstAuthor="Yu" journalOrPublisher="Chenia" pagination="58-62" refId="B146" refString="Yu NN, Gradstein SR, Narengaowa (2020) Thysananthus weiweianus N. - N. Yu &amp; Gradst. (Jungermanniopsida: Lejeuneaceae), a new fossil species in Dominican amber. Chenia 14: 5862." title="Thysananthus weiweianus N. - N. Yu &amp; Gradst. (Jungermanniopsida: Lejeuneaceae), a new fossil species in Dominican amber." volume="14" year="2020">Yu et al. 2020</bibRefCitation>
).
<taxonomicName id="1427DD4F40BDAE909CC48348FB9312EB" class="Jungermanniopsida" family="Lejeuneaceae" genus="Thysananthus" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="auriculatus">
<emphasis id="29679D9DDD1C4DC6CC36BF98557EBF18" italics="true">Thysananthus auriculatus</emphasis>
</taxonomicName>
can be differentiated from
<taxonomicName id="1AD46197ECD394BF4192BD1017C5B948" authorityName="K. Feldberg, Gradst., Schaf. - Verw., Mamontov" authorityYear="2025" class="Jungermanniopsida" family="Lejeuneaceae" genus="Thysananthus" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="patrickmuelleri">
<emphasis id="6FE12DBFF15F684E845AC02B0D372FA3" italics="true">T. patrickmuelleri</emphasis>
</taxonomicName>
by the less elongate leaf lobes, the more strongly incurved postical leaf margin, the oblique apical free margin of the lobule which continues into the postical margin of the lobe, and the either small or absent apical lobule tooth (
<bibRefCitation id="A632EE435AB66724C2BA8DCC573AD1C1" author="Gradstein" firstAuthor="Gradstein" journalOrPublisher="Nova Hedwigia" pagination="353-374" refId="B31" refString="Gradstein SR (1993) New fossil Hepaticae preserved in amber of the Dominican Republic. Nova Hedwigia 57: 353374." title="New fossil Hepaticae preserved in amber of the Dominican Republic." volume="57" year="1993">Gradstein 1993</bibRefCitation>
, fig. 7;
<bibRefCitation id="D2512E0C04596064F650469ADD164BB7" DOI="10.1127/0029-5035/2014/0206" author="Sukkharak and Gradstein" firstAuthor="Sukkharak" journalOrPublisher="Nova Hedwigia" pagination="279-345" refId="B130" refString="Sukkharak P, Gradstein SR (2014) A taxonomic revision of the genus Mastigolejeunea (Marchantiophyta: Lejeuneaceae). Nova Hedwigia 99: 279345. https://doi.org/10.1127/0029-5035/2014/0206" title="A taxonomic revision of the genus Mastigolejeunea (Marchantiophyta: Lejeuneaceae)." volume="99" year="2014">Sukkharak and Gradstein 2014</bibRefCitation>
, figs 4, 5).
<taxonomicName id="9C55A086F165484B846151A76D9BA13E" class="Jungermanniopsida" family="Lejeuneaceae" genus="Thysananthus" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="bidentulus">
<emphasis id="48AF5AF74E9E896CF84C441EE5DC4A5E" italics="true">Thysananthus bidentulus</emphasis>
</taxonomicName>
is more similar to
<taxonomicName id="ABD79BA86B51B45AA96C9FBAC289E178" authorityName="K. Feldberg, Gradst., Schaf. - Verw., Mamontov" authorityYear="2025" class="Jungermanniopsida" family="Lejeuneaceae" genus="Thysananthus" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="patrickmuelleri">
<emphasis id="108156507B86C94F7CCC5A03E7F7A8CC" italics="true">T. patrickmuelleri</emphasis>
</taxonomicName>
in its general habit, but differs in having oblong lobules with oblique apices, which continue into the postical margin of the lobe, and the presence of two one-celled lobule teeth (
<bibRefCitation id="0EF64C84843DE819B3864BF6F61BD20E" author="Gradstein" firstAuthor="Gradstein" journalOrPublisher="Nova Hedwigia" pagination="353-374" refId="B31" refString="Gradstein SR (1993) New fossil Hepaticae preserved in amber of the Dominican Republic. Nova Hedwigia 57: 353374." title="New fossil Hepaticae preserved in amber of the Dominican Republic." volume="57" year="1993">Gradstein 1993</bibRefCitation>
, fig. 8).
<taxonomicName id="616CA9716562C84EB0E68DEDAC28DCBC" class="Jungermanniopsida" family="Lejeuneaceae" genus="Thysananthus" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="weiweianus">
<emphasis id="3CE589FD531AEC5853B919BD85A1C9C7" italics="true">Thysananthus weiweianus</emphasis>
</taxonomicName>
differs from the new fossil species in having larger, often confluent trigones, a very small or absent lobule tooth, and more elongate underleaves with plane lateral margins (
<bibRefCitation id="55F54C820E04182CD293471CDCF37DD6" author="Yu" etAl="et al." firstAuthor="Yu" journalOrPublisher="Chenia" pagination="58-62" refId="B146" refString="Yu NN, Gradstein SR, Narengaowa (2020) Thysananthus weiweianus N. - N. Yu &amp; Gradst. (Jungermanniopsida: Lejeuneaceae), a new fossil species in Dominican amber. Chenia 14: 5862." title="Thysananthus weiweianus N. - N. Yu &amp; Gradst. (Jungermanniopsida: Lejeuneaceae), a new fossil species in Dominican amber." volume="14" year="2020">Yu et al. 2020</bibRefCitation>
, figs 1, 2). Another amber deposit that includes
<taxonomicName id="5B888410AE5D7021D8B58BD020AECF79" class="Jungermanniopsida" family="Lejeuneaceae" genus="Thysananthus" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="genus">
<emphasis id="E6B5F941A3688622C89A805F24B4FECA" italics="true">Thysananthus</emphasis>
</taxonomicName>
is Miocene Ethiopian Shewa amber (
<bibRefCitation id="8DE3CF68CFD5A66E6604042A6FF8C385" DOI="10.1111/jse.12796" author="Bouju" etAl="et al." firstAuthor="Bouju" journalOrPublisher="Journal of Systematics and Evolution" pagination="932-954" refId="B9" refString="Bouju V, Feldberg K, Kaasalainen U, Schäfer-Verwimp A, Hedenäs L, Buck WR, Wang B, Perrichot V, Schmidt AR (2021) Miocene Ethiopian amber: A new source of fossil cryptogams. Journal of Systematics and Evolution 60: 932954. https://doi.org/10.1111/jse.12796" title="Miocene Ethiopian amber: A new source of fossil cryptogams." volume="60" year="2021">Bouju et al. 2021</bibRefCitation>
, fig. 3).
<taxonomicName id="F455BC3D7114822D35A6EDC2CB858413" authority="Bouju et al." authorityName="Bouju" class="Jungermanniopsida" family="Lejeuneaceae" genus="Thysananthus" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="aethiopicus">
<emphasis id="DC9F1357F186FDC18F7B08B1B43FF56B" italics="true">Thysananthus aethiopicus</emphasis>
Bouju et al.
</taxonomicName>
differs from the new Mexican fossil in having lobes with more strongly incurved postical margins and occasionally apiculate apices, as well as lobules with a very coarse apical tooth and sometimes a second tooth.
</paragraph>
<paragraph id="80EAF51CEA2E60AF8F1D44147608CEDA">
A specimen similar to
<taxonomicName id="FFEBA0FAC9105A369D8A0F3E2EB3620E" authorityName="K. Feldberg, Gradst., Schaf. - Verw., Mamontov" authorityYear="2025" class="Jungermanniopsida" family="Lejeuneaceae" genus="Thysananthus" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="patrickmuelleri">
<emphasis id="64BCC5101213AC7CF4B386EA2178AE20" italics="true">T. patrickmuelleri</emphasis>
</taxonomicName>
in its general habit has already been found in Mexican amber but was only identified as belonging to
<taxonomicName id="DE7F81C846493254EB06ABFDE55CA57A" class="Jungermanniopsida" family="Lejeuneaceae" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="subFamily" subFamily="Ptychanthoideae">Ptychanthoideae</taxonomicName>
and possibly to
<taxonomicName id="192689A293B3EE9878AF5C1CBA680682" class="Jungermanniopsida" family="Lejeuneaceae" genus="Mastigolejeunea" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="genus">
<emphasis id="B9462475C02E565A22B40C8ADE976F97" italics="true">Mastigolejeunea</emphasis>
</taxonomicName>
(Fig.
<figureCitation id="681CD9533AD3FE34ED360664FEDE8376" captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. A E. Thysananthus patrickmuelleri sp. nov. (A E, Patrick Müller amber collection, MEX 71) and a representative of Ptychanthoideae (F, Stuttgart State Museum of Natural History, Mx 357) from Mexican amber. A. Branched gametophyte of Thysananthus patrickmuelleri in dorso-lateral view; B. Left shoot with erect-spreading leaves similar to MEX 70 and right shoot with spreading leaves similar to holotype GZG. BST. 22086; C. Lower part with several flagelliform branches, the one on the left becoming main shoot-like; D. Flagelliform branch in ventral view; E. Detail of the main shoot in ventral view; F. Ptychanthoideae sp., gametophyte in ventral view." figureDoi="10.3897/fr.28.137758.figure6" httpUri="https://binary.pensoft.net/fig/1264818">6 F</figureCitation>
;
<bibRefCitation id="9DD50FBE2198BD781D8E284BD8EC5CD9" DOI="10.7751/telopea20148083" author="Scheben" etAl="et al." firstAuthor="Scheben" journalOrPublisher="Telopea" pagination="355-361" refId="B115" refString="Scheben A, Schmidt AR, Schäfer-Verwimp A, Solórzano Kraemer MM, Heinrichs J (2014) The first ptychanthoid Lejeuneaceae in Miocene Mexican amber. Telopea 17: 355361. https://doi.org/10.7751/telopea20148083" title="The first ptychanthoid Lejeuneaceae in Miocene Mexican amber." volume="17" year="2014">Scheben et al. 2014</bibRefCitation>
). Many important diagnostic characters are not visible in this specimen, especially the walls of the lobe cells, which are crucial for correctly identifying the genus, and the structure of the lobules.
</paragraph>
<paragraph id="5B99B14615C96225058C0CC52A69F226">
Compared to the extant species of
<taxonomicName id="EE547E4403360E15A1C2CA359619B7DE" class="Jungermanniopsida" family="Lejeuneaceae" genus="Thysananthus" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="genus">
<emphasis id="1BA1AF1D0F3DD18A7C8C01164F63B1BF" italics="true">Thysananthus</emphasis>
</taxonomicName>
, the new fossil is very similar to the neotropical
<taxonomicName id="BC5DA9C60CD4CD00D1C82DFE83FF4178" authority="(Spruce) Sukkharak &amp; Gradst." authorityName="Sukkharak &amp; Gradst." baseAuthorityName="Spruce" class="Jungermanniopsida" family="Lejeuneaceae" genus="Thysananthus" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="plicatiflorus">
<emphasis id="1004EBDF4188BF9645A8653ACBFE935C" italics="true">T. plicatiflorus</emphasis>
(Spruce) Sukkharak &amp; Gradst.
</taxonomicName>
, a species widely distributed in the rainforests of northern South America (
<bibRefCitation id="A6B0DF5E576E56B2AC2836DCB5D8ED73" author="Gradstein" firstAuthor="Gradstein" journalOrPublisher="Flora Neotropica Monograph" pagination="1-225" refId="B32" refString="Gradstein SR (1994) Lejeuneaceae: Ptychantheae. Brachiolejeuneae. Flora Neotropica Monograph 62: 1225." title="Lejeuneaceae: Ptychantheae. Brachiolejeuneae." volume="62" year="1994">Gradstein 1994</bibRefCitation>
,
<bibRefCitation id="BD1F50D758B394DD39D66CE3EF21A5AB" DOI="10.1007/978-3-030-49450-6" author="Gradstein" firstAuthor="Gradstein" refId="B36" refString="Gradstein SR (2021) The Liverworts and Hornworts of Colombia and Ecuador. Memoirs of the New York Botanical Garden 121: i xxv, 1723. https://doi.org/10.1007/978-3-030-49450-6" year="2021">2021</bibRefCitation>
;
<bibRefCitation id="8B9882D31F09527009768FCB8FF2F017" DOI="10.1127/0029-5035/2014/0206" author="Sukkharak and Gradstein" firstAuthor="Sukkharak" journalOrPublisher="Nova Hedwigia" pagination="279-345" refId="B130" refString="Sukkharak P, Gradstein SR (2014) A taxonomic revision of the genus Mastigolejeunea (Marchantiophyta: Lejeuneaceae). Nova Hedwigia 99: 279345. https://doi.org/10.1127/0029-5035/2014/0206" title="A taxonomic revision of the genus Mastigolejeunea (Marchantiophyta: Lejeuneaceae)." volume="99" year="2014">Sukkharak and Gradstein 2014</bibRefCitation>
, fig. 21, as
<taxonomicName id="102B309D38B055FC5F6C54B1E4B82779" authority="(Spruce) Steph." authorityName="Steph." baseAuthorityName="Spruce" class="Jungermanniopsida" family="Lejeuneaceae" genus="Mastigolejeunea" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="plicatiflora">
<emphasis id="62BFA2BD585842B56B3C59DE1B25636E" italics="true">Mastigolejeunea plicatiflora</emphasis>
(Spruce) Steph.
</taxonomicName>
). Both species share elongate, ovate-oblong leaves with rounded apices and a plane postical margin, lobules with a single short tooth and a truncate apical free margin that terminates at the end of the keel, as well as auriculate underleaves. The two species differ markedly, however, in the orientation of the underleaves, which are very flat and have a plane apex in
<taxonomicName id="E8CD063E684B97A5AFDF16EF4FA842EF" class="Jungermanniopsida" family="Lejeuneaceae" genus="Thysananthus" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="plicatiflorus">
<emphasis id="DE44CCE2C157172007ECDC698D3B92BD" italics="true">T. plicatiflorus</emphasis>
</taxonomicName>
while being somewhat squarrose with a distinctly recurved apex in
<taxonomicName id="3B7DE3D751A35557B06FEAA4C1F2E846" authorityName="K. Feldberg, Gradst., Schaf. - Verw., Mamontov" authorityYear="2025" class="Jungermanniopsida" family="Lejeuneaceae" genus="Thysananthus" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="patrickmuelleri">
<emphasis id="8D569A169CC3CB70A911B0F1B2F452BB" italics="true">T. patrickmuelleri</emphasis>
</taxonomicName>
(apex only occasionally plane on branch leaves). Another quite similar species is the Australasian
<taxonomicName id="51F8CA713E6319E41761EC282431B691" authority="(Lehm. &amp; Lindenb.) Sukkharak &amp; Gradst." authorityName="Sukkharak &amp; Gradst." baseAuthorityName="Lehm. &amp; Lindenb." class="Jungermanniopsida" family="Lejeuneaceae" genus="Thysananthus" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="ligulatus">
<emphasis id="BCCD255320D83C195C7793CE008A44BE" italics="true">T. ligulatus</emphasis>
(Lehm. &amp; Lindenb.) Sukkharak &amp; Gradst.
</taxonomicName>
(
<bibRefCitation id="6FDE3C9C2CE991AA665C0C56A932CD31" DOI="10.1127/0029-5035/2014/0206" author="Sukkharak and Gradstein" firstAuthor="Sukkharak" journalOrPublisher="Nova Hedwigia" pagination="279-345" refId="B130" refString="Sukkharak P, Gradstein SR (2014) A taxonomic revision of the genus Mastigolejeunea (Marchantiophyta: Lejeuneaceae). Nova Hedwigia 99: 279345. https://doi.org/10.1127/0029-5035/2014/0206" title="A taxonomic revision of the genus Mastigolejeunea (Marchantiophyta: Lejeuneaceae)." volume="99" year="2014">Sukkharak and Gradstein 2014</bibRefCitation>
, figs 18, 19, as
<taxonomicName id="6ACEC8BDCFA989477089A82392E8A19B" authority="(Lehm. &amp; Lindenb.) Schiffn." authorityName="Schiffn." baseAuthorityName="Lehm. &amp; Lindenb." class="Jungermanniopsida" family="Lejeuneaceae" genus="Mastigolejeunea" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="ligulata">
<emphasis id="5DDD4697F5F48C38006022C01BF1145A" italics="true">Mastigolejeunea ligulata</emphasis>
(Lehm. &amp; Lindenb.) Schiffn.
</taxonomicName>
), which also has rather elongated lobes with rounded apices, plane margins, and auriculate underleaves. However, it can be differentiated by the lobules being smaller in relation to the lobes (0.20.3 times the lobe length vs. 0.30.4) and underleaves that are often longer than wide, spathulate, and not overlapping the stem as much as those of
<taxonomicName id="C6279F998138E81C6AEBFD3C0070C9A1" authorityName="K. Feldberg, Gradst., Schaf. - Verw., Mamontov" authorityYear="2025" class="Jungermanniopsida" family="Lejeuneaceae" genus="Thysananthus" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="species" species="patrickmuelleri">
<emphasis id="5286E496DA527EE5992F9532B4BE1F27" italics="true">T. patrickmuelleri</emphasis>
</taxonomicName>
(34 times wider than the stem vs. 36). Since the new fossil shows significant morphological differences to all known extinct and extant species of
<taxonomicName id="23CCE8A6184E2E4F84F8BDF526812198" class="Jungermanniopsida" family="Lejeuneaceae" genus="Thysananthus" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="genus">
<emphasis id="0FA61DA3A4E73E00BD71FB4631B7D814" italics="true">Thysananthus</emphasis>
</taxonomicName>
, it is described as a new species.
</paragraph>
</subSubSection>
</treatment>
</document>

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<mods:detail id="F56CA8714C57B2FC7A832855E033F9A1" type="volume">
<mods:number id="0DBC68F221BD9395B88AF80C94AEA206">28</mods:number>
</mods:detail>
<mods:detail id="44656DAF3255050CB27F1A376ADDB0CB" type="issue">
<mods:number id="AFD215193D553424A3CDE15E8FE8CC9B">1</mods:number>
</mods:detail>
<mods:extent id="9F8A35A6F50618A30CD1D68ADECD2E14" unit="page">
<mods:start id="5FA06F421A437491569B74265E1B1B2B">79</mods:start>
<mods:end id="26E718C6FF16A3E3AB016ECE14A2AF2E">102</mods:end>
</mods:extent>
</mods:part>
</mods:relatedItem>
<mods:classification id="BA05BF0FC5CDA34ADA21F24C3EB7C7B9">journal article</mods:classification>
<mods:identifier id="BDAD76A33C2D121B5ADFF855DBADE043" type="DOI">10.3897/fr.28.137758</mods:identifier>
<mods:identifier id="EAA5B41BC49DB88DEFF945CC019E7F0E" type="ZooBank">A10C1262-F859-4004-A83C-C8F032C1156A</mods:identifier>
</mods:mods>
<treatment id="BA36D971E7925D309496C443A93304A0" ID-arpha="BA36D971-E792-5D30-9496-C443A93304A0" LSID="urn:lsid:plazi:treatment:BA36D971E7925D309496C443A93304A0" httpUri="http://treatment.plazi.org/id/BA36D971E7925D309496C443A93304A0" scope_class="Lecanoromycetes" scope_order="Lecanorales" scope_phylum="Ascomycota">
<subSubSection id="E785637DA53F1CCC928BBFEDA7682758" type="nomenclature">
<paragraph id="26665775B5937547C1F4E7B85247AB7E">
<taxonomicName id="8243C4017C6347B2A49ACED605A59858" class="Lecanoromycetes" family="Parmeliaceae" genus="Parmotrema" kingdom="Fungi" order="Lecanorales" phylum="Ascomycota" rank="species" species="undetermined-1">
<emphasis id="2938BAB33975162BC574D6FD9B8F97F1" italics="true">Parmotrema</emphasis>
specimen 1
</taxonomicName>
</paragraph>
</subSubSection>
<subSubSection id="SECID0EO4AI" type="Specimen">
<paragraph id="60F423C0A704974221CB78E1BC19E930">
<heading id="D856CD1D1E674380956654BDD04CC83F" reason="title">Specimen.</heading>
</paragraph>
<paragraph id="6CD3A381524E96EA56B03E8ECF2AD3B5">
<named-content id="F6B6633DB3B4484439835A0EB27438BD" content-type="institution" xlink_href="http://grbio.org/institution/staatliches-museum-fuer-naturkund-stuttgart" xlink_type="simple">
Stuttgart State Museum of Natural History,
<collectingCountry id="8883051FDC9792F375828778F5CFB079" name="Germany">Germany</collectingCountry>
</named-content>
(
<named-content id="6DE452DEB3650281DBA818A5DC94B102" content-type="dwc:institutional_code" xlink_href="http://grbio.org/institution/staatliches-museum-fuer-naturkund-stuttgart" xlink_title="Stuttgart State Museum of Natural History, Germany">
<collectionCode id="84AC37D94F7C5D626856282B8B6F9EF5" country="Germany" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34840" lsid="urn:lsid:biocol.org:col:34840" name="Staatliches Museum fuer Naturkund Stuttgart" type="">SMNS</collectionCode>
</named-content>
),
<specimenCode id="BABCA66694F641C0894E7EFE609C61ED">
<named-content id="80C291709451530C631F8B9357B090B5" content-type="dwc:institutional_code" xlink_href="http://grbio.org/institution/staatliches-museum-fuer-naturkund-stuttgart" xlink_title="Stuttgart State Museum of Natural History, Germany">SMNS</named-content>
-DO-4928-M
</specimenCode>
; syninclusion unidentified leafy liverwort of the order
<taxonomicName id="F7FAD0AEAE6C9CB56BA9D91BC00CCD7A" authorityName="Schljakov" authorityYear="1972" class="Jungermanniopsida" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="order">Porellales</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection id="SECID0ED5AI" type="Age and stratigraphic level">
<paragraph id="35BD7792AAAA39293E08705139CD1FF5">
<heading id="47B0FC788969DEE18978F02738C22D59" reason="title">Age and stratigraphic level.</heading>
</paragraph>
<paragraph id="1B5135D5C19534DBD44218E697CC00A8">
1520 Ma, Langhian Burdigalian (early to middle Miocene), La Toca Formation,
<collectingCountry id="390409BF67BF79A3E9B126E24B87C44A" name="Dominican Republic">Dominican Republic</collectingCountry>
.
</paragraph>
</subSubSection>
<subSubSection id="SECID0ES5AI" type="description">
<paragraph id="4E90355F73B03341D58400B4FB7E2C89">
<heading id="C229851B1A1FAB0BC5210BA0722AED28" reason="title">Description.</heading>
</paragraph>
<paragraph id="04BBDA07C8B07C79FDDFEF917212E5CD">
Lichen fragment approximately 5.4 ×
<quantity id="FF7D90C86FD8C0F8F891748EA5E3E407" metricMagnitude="-3" metricUnit="m" metricValue="2.6" unit="mm" value="2.6">2.6 mm</quantity>
in diameter.
<emphasis id="2C78785C4902CCC7E4FF3BC79DDE06D1" bold="true" italics="true">Thallus</emphasis>
foliose, lobate.
<emphasis id="FA8A04A3E1C9A0214C74394B70AE70B3" bold="true" italics="true">Lobes</emphasis>
robust, flat, and linear,
<quantity id="100CD74844E2BAEA13E19C4AF913E332" metricMagnitude="-4" metricUnit="m" metricValue="6.5" metricValueMax="9.0" metricValueMin="4.0" unit="mm" value="0.65" valueMax="0.9" valueMin="0.4">0.40.9 mm</quantity>
wide, lobe apices truncate (Fig.
<figureCitation id="BE6BD3E0AB65A9E8CBAB4180D71D1C43" captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. Parmotrema fossils from Dominican amber. A-C. Parmotrema specimen 1 (SMNS - DO- 4928 - M); A. Parmotrema inclusion with a bryophyte (leafy liverwort, Porellales); B. Lobes seen from above showing the truncate lobe apices, dark lobe margins, and long marginal cilia. The lighter spots visible on the upper surface of the thallus are caused by slight depressions and their brighter color derives from air-filled spaces between the thallus surface and amber. These depressions seem to correspond to the attachment points of the rhizines on the lower surface; C. Close-up of the slightly uneven upper surface and finger-like isidia. D-E. Parmotrema specimen 2 (SMNS - DO- 4929 - M); D. Close-up of the upper surface with abundant pycnidia; E. General view of the inclusion, showing the very long, branching marginal cilia." figureDoi="10.3897/fr.28.137758.figure7" httpUri="https://binary.pensoft.net/fig/1264819">7 B</figureCitation>
).
<emphasis id="773BA7EF7C055B83BA079D9CBF817B16" bold="true" italics="true">Upper surface</emphasis>
slightly uneven, brown, with prominent dark margins (Fig.
<figureCitation id="8959D1C753A9B23A0E9C1C0E77F32BD5" captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. Parmotrema fossils from Dominican amber. A-C. Parmotrema specimen 1 (SMNS - DO- 4928 - M); A. Parmotrema inclusion with a bryophyte (leafy liverwort, Porellales); B. Lobes seen from above showing the truncate lobe apices, dark lobe margins, and long marginal cilia. The lighter spots visible on the upper surface of the thallus are caused by slight depressions and their brighter color derives from air-filled spaces between the thallus surface and amber. These depressions seem to correspond to the attachment points of the rhizines on the lower surface; C. Close-up of the slightly uneven upper surface and finger-like isidia. D-E. Parmotrema specimen 2 (SMNS - DO- 4929 - M); D. Close-up of the upper surface with abundant pycnidia; E. General view of the inclusion, showing the very long, branching marginal cilia." figureDoi="10.3897/fr.28.137758.figure7" httpUri="https://binary.pensoft.net/fig/1264819">7 C</figureCitation>
).
<emphasis id="6C38147368757ECD43C64A5456AC819A" bold="true" italics="true">Medulla</emphasis>
not visible.
<emphasis id="797E5941C19A81BC6544242CDF4EA52F" bold="true" italics="true">Lower cortex</emphasis>
dark.
<emphasis id="74C6FD83F512590CDCCB6967D3FB27A1" bold="true" italics="true">Marginal cilia</emphasis>
long (up to at least
<quantity id="29D9A1DD0E65927001A03965318497B2" metricMagnitude="-4" metricUnit="m" metricValue="8.0" unit="mm" value="0.8">0.8 mm</quantity>
), dark, thick and tapered (Fig.
<figureCitation id="0C961D23E6783771FEF110530063DCC3" captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. Parmotrema fossils from Dominican amber. A-C. Parmotrema specimen 1 (SMNS - DO- 4928 - M); A. Parmotrema inclusion with a bryophyte (leafy liverwort, Porellales); B. Lobes seen from above showing the truncate lobe apices, dark lobe margins, and long marginal cilia. The lighter spots visible on the upper surface of the thallus are caused by slight depressions and their brighter color derives from air-filled spaces between the thallus surface and amber. These depressions seem to correspond to the attachment points of the rhizines on the lower surface; C. Close-up of the slightly uneven upper surface and finger-like isidia. D-E. Parmotrema specimen 2 (SMNS - DO- 4929 - M); D. Close-up of the upper surface with abundant pycnidia; E. General view of the inclusion, showing the very long, branching marginal cilia." figureDoi="10.3897/fr.28.137758.figure7" httpUri="https://binary.pensoft.net/fig/1264819">7 B, C</figureCitation>
).
<emphasis id="D9107044D85AD685AB0B47C522D93138" bold="true" italics="true">Rhizines</emphasis>
dark, shorter than cilia.
<emphasis id="E581D363C3EB0A9CAB46B5E60DAC6E99" bold="true" italics="true">Isidia</emphasis>
abundant, laminal, finger-shaped, up to 150 µm long (Fig.
<figureCitation id="B964C6520EB3B5F47C1450A444411387" captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. Parmotrema fossils from Dominican amber. A-C. Parmotrema specimen 1 (SMNS - DO- 4928 - M); A. Parmotrema inclusion with a bryophyte (leafy liverwort, Porellales); B. Lobes seen from above showing the truncate lobe apices, dark lobe margins, and long marginal cilia. The lighter spots visible on the upper surface of the thallus are caused by slight depressions and their brighter color derives from air-filled spaces between the thallus surface and amber. These depressions seem to correspond to the attachment points of the rhizines on the lower surface; C. Close-up of the slightly uneven upper surface and finger-like isidia. D-E. Parmotrema specimen 2 (SMNS - DO- 4929 - M); D. Close-up of the upper surface with abundant pycnidia; E. General view of the inclusion, showing the very long, branching marginal cilia." figureDoi="10.3897/fr.28.137758.figure7" httpUri="https://binary.pensoft.net/fig/1264819">7 C</figureCitation>
). Apothecia, soredia, or pycnidia not present.
</paragraph>
<caption id="8DC32B40A205F204812FABB7D4F18A7C" ID-DOI="10.3897/fr.28.137758.figure7" ID-arpha="8F3E3463-81A0-5CA4-B8E2-94C369ED0236" httpUri="https://binary.pensoft.net/fig/1264819" startId="F7">
<paragraph id="9BDB1E64E448F2A7CE45003E283F8A7A">
<label id="AC42277A54011ECC061EC765017161BA">Figure 7.</label>
</paragraph>
<paragraph id="D56C00E63D036546EF0DCB22C74FAF3A">
<taxonomicName id="39EFACF918C9782894792859FA9F05A5" class="Lecanoromycetes" family="Parmeliaceae" genus="Parmotrema" kingdom="Fungi" order="Lecanorales" phylum="Ascomycota" rank="genus">
<emphasis id="F580B834DE5134808B7B56027BB0171B" italics="true">Parmotrema</emphasis>
</taxonomicName>
fossils from Dominican amber.
<emphasis id="3D2F4C7D8554C3925A4424DB03A8260B" bold="true">A-C.</emphasis>
<taxonomicName id="AE9076144CCF5AD371A5C0EC567A30B9" class="Lecanoromycetes" family="Parmeliaceae" genus="Parmotrema" kingdom="Fungi" order="Lecanorales" phylum="Ascomycota" rank="genus">
<emphasis id="0D2E3209F743AD22C773C2FE3B419AD2" italics="true">Parmotrema</emphasis>
</taxonomicName>
specimen 1 (
<specimenCode id="809A8D95F70DBA7FC969C5B8F31ECE6E">
<named-content id="1F843676979506B6BF9F82A8F27DCF66" content-type="dwc:institutional_code" xlink_href="http://grbio.org/institution/staatliches-museum-fuer-naturkund-stuttgart" xlink_title="Stuttgart State Museum of Natural History, Germany">SMNS</named-content>
-DO-4928-M
</specimenCode>
);
<emphasis id="53B2A5784E17761A6E7BAC80E2BED5D9" bold="true">A.</emphasis>
<taxonomicName id="7E39EA8618A44E12DD969BCA94E68620" class="Lecanoromycetes" family="Parmeliaceae" genus="Parmotrema" kingdom="Fungi" order="Lecanorales" phylum="Ascomycota" rank="genus">
<emphasis id="D10CBE1FD8EC94482EE4A7E0CC0944B9" italics="true">Parmotrema</emphasis>
</taxonomicName>
inclusion with a bryophyte (leafy liverwort,
<taxonomicName id="EF357C8EA15CB886CBEFCD0FF49EAAA2" authorityName="Schljakov" authorityYear="1972" class="Jungermanniopsida" kingdom="Plantae" order="Porellales" phylum="Marchantiophyta" rank="order">Porellales</taxonomicName>
);
<emphasis id="DBBD6F1493C19D54F40759D9950DCA1C" bold="true">B.</emphasis>
Lobes seen from above showing the truncate lobe apices, dark lobe margins, and long marginal cilia. The lighter spots visible on the upper surface of the thallus are caused by slight depressions and their brighter color derives from air-filled spaces between the thallus surface and amber. These depressions seem to correspond to the attachment points of the rhizines on the lower surface;
<emphasis id="6C42584925CA49CE99B7D06F8953B06D" bold="true">C.</emphasis>
Close-up of the slightly uneven upper surface and finger-like isidia.
<emphasis id="DE26ABBF8448170B7F6CF5FC5C865400" bold="true">D-E.</emphasis>
<taxonomicName id="EF1D0E2BF15EEDAE96A935192F8D97B0" class="Lecanoromycetes" family="Parmeliaceae" genus="Parmotrema" kingdom="Fungi" order="Lecanorales" phylum="Ascomycota" rank="genus">
<emphasis id="17FC989AAD92EBB3FAA6B738F26A660C" italics="true">Parmotrema</emphasis>
</taxonomicName>
specimen 2 (
<specimenCode id="8758974AD4F1784B1F100C3B4F2DD72D">
<named-content id="B9590B966A2CEEF6F6E3DF80310E15DB" content-type="dwc:institutional_code" xlink_href="http://grbio.org/institution/staatliches-museum-fuer-naturkund-stuttgart" xlink_title="Stuttgart State Museum of Natural History, Germany">SMNS</named-content>
-DO-4929-M
</specimenCode>
);
<emphasis id="E99A65D95844A20E819E02A028D2B921" bold="true">D.</emphasis>
Close-up of the upper surface with abundant pycnidia;
<emphasis id="A56036D6A0C43F98DFE1E98049484E68" bold="true">E.</emphasis>
General view of the inclusion, showing the very long, branching marginal cilia.
</paragraph>
</caption>
</subSubSection>
<subSubSection id="SECID0EFDBI" type="discussion">
<paragraph id="B296FE671B072ED0CBDDEFFDE054EB60">
<heading id="AAEF4ACBCA5B002CAB2299F4603AE0CF" reason="title">Discussion.</heading>
</paragraph>
<paragraph id="8F7E23D8A0F132E5B5A45F8B9B83B416">
The general habit and long, thick, and tapered marginal cilia identify the specimen as
<taxonomicName id="7E4AEC5FA42F4BB2273E5F2B714BED4C" class="Lecanoromycetes" family="Parmeliaceae" genus="Parmotrema" kingdom="Fungi" order="Lecanorales" phylum="Ascomycota" rank="genus">
<emphasis id="25159C72C9CB52CE5C89F01079A3BBC3" italics="true">Parmotrema</emphasis>
</taxonomicName>
.
<taxonomicName id="33D2B8851563F0F48448AA2363905DEE" class="Lecanoromycetes" family="Parmeliaceae" genus="Parmotrema" kingdom="Fungi" order="Lecanorales" phylum="Ascomycota" rank="genus">
<emphasis id="ECAFC71BD6E96A59A7A9E410DE1FF051" italics="true">Parmotrema</emphasis>
</taxonomicName>
is an extant genus with approximately 300 species mainly distributed in tropical and subtropical regions, especially in the Pacific Islands and South America (
<bibRefCitation id="69476BA8AE292F5125EFF1550C0B199A" DOI="10.1111/j.1756-1051.2012.00008.x" author="Thell" etAl="et al." firstAuthor="Thell" refId="B133" refString="Thell A, Crespo A, Divakar PK, Kärnefelt I, Leavitt SD, Lumbsch HT, Seaward MRD (2012) A review of the lichen family Parmeliaceae history, phylogeny and current taxonomy. Nordic Journal of Botany 30, 641664. https://doi.org/10.1111/j.1756-1051.2012.00008.x" year="2012">Thell et al. 2012</bibRefCitation>
). It is part of the parmelioid crown group of the
<taxonomicName id="4A413A4A3E3DDAE672897DB6192F63FE" authority="(Pizarro et al. 2018)" baseAuthorityName="Pizarro" baseAuthorityYear="2018" class="Lecanoromycetes" family="Parmeliaceae" kingdom="Fungi" order="Lecanorales" phylum="Ascomycota" rank="family">
Parmeliaceae (
<bibRefCitation id="541183E81A49D54E290930451FF417B8" DOI="10.1007/s13225-018-0407-7" author="Pizarro" etAl="et al." firstAuthor="Pizarro" journalOrPublisher="Fungal Diversity" pagination="31-41" refId="B99" refString="Pizarro D, Divakar PK, Grewe F, Leavitt SD, Huang J-P, Dal Grande F, Schmitt I, Wedin M, Crespo A, Lumbsch HT (2018) Phylogenomic analysis of 2556 single-copy protein-coding genes resolves most evolutionary relationships for the major clades in the most diverse group of lichen-forming fungi. Fungal Diversity 92: 3141. https://doi.org/10.1007/s13225-018-0407-7" title="Phylogenomic analysis of 2556 single-copy protein-coding genes resolves most evolutionary relationships for the major clades in the most diverse group of lichen-forming fungi." volume="92" year="2018">Pizarro et al. 2018</bibRefCitation>
)
</taxonomicName>
where apothecial and conidial characters, growth form, cortical and medullar chemistry, and presence, absence and / or type of cell-wall polysaccharides, marginal cilia, rhizines, and surface features, like epicortex and pseudocyphellae, have been used to separate genera (
<bibRefCitation id="753ECF9FCC6A21F1B341BF2B505CA161" DOI="10.1017/S0024282911000570" author="Crespo" etAl="et al." firstAuthor="Crespo" journalOrPublisher="The Lichenologist" pagination="511-535" refId="B13" refString="Crespo A, Divakar PK, Hawksworth DL (2011) Generic concepts in parmelioid lichens, and the phylogenetic value of characters used in their circumscription. The Lichenologist 43: 511535. https://doi.org/10.1017/S0024282911000570" title="Generic concepts in parmelioid lichens, and the phylogenetic value of characters used in their circumscription." volume="43" year="2011">Crespo et al. 2011</bibRefCitation>
). Like in many parmelioid genera, molecular phylogenetics have repeatedly shaped the generic boundaries of
<taxonomicName id="1B57208D01526515FE4915726ECB0A66" class="Lecanoromycetes" family="Parmeliaceae" genus="Parmotrema" kingdom="Fungi" order="Lecanorales" phylum="Ascomycota" rank="genus">
<emphasis id="7D02E72D1D6C0C0E7F91EFA4B0733DF9" italics="true">Parmotrema</emphasis>
</taxonomicName>
(e. g.,
<bibRefCitation id="5872E17090DB1B0E409E35128AD9C56F" DOI="10.1080/15572536.2006.11832848" author="Blanco" etAl="et al." firstAuthor="Blanco" journalOrPublisher="Mycologia" pagination="150-159" refId="B7" refString="Blanco O, Crespo A, Divakar PK, Elix JA, Lumbsch HT (2005) Molecular phylogeny of parmotremoid lichens (Ascomycota, Parmeliaceae). Mycologia 97: 150159. https://doi.org/10.1080/15572536.2006.11832848" title="Molecular phylogeny of parmotremoid lichens (Ascomycota, Parmeliaceae)." volume="97" year="2005">Blanco et al. 2005</bibRefCitation>
;
<bibRefCitation id="9262C8F6EB2A1918E31B65F3EA229552" DOI="10.1017/S0024282904014586" author="Divakar" etAl="et al." firstAuthor="Divakar" journalOrPublisher="The Lichenologist" pagination="55-65" refId="B15" refString="Divakar PK, Blanco O, Hawksworth DL, Crespo A (2005) Molecular phylogenetic studies on the Parmotrema reticulatum (syn. Rimelia reticulata) complex, including the confirmation of P. pseudoreticulatum as a distinct species. The Lichenologist 37: 5565. https://doi.org/10.1017/S0024282904014586" title="Molecular phylogenetic studies on the Parmotrema reticulatum (syn. Rimelia reticulata) complex, including the confirmation of P. pseudoreticulatum as a distinct species." volume="37" year="2005">Divakar et al. 2005</bibRefCitation>
,
<bibRefCitation id="6B65896A5D0F85E16EC9BCFC350E7B77" DOI="10.1007/s13225-017-0379-z" author="Divakar" firstAuthor="Divakar" journalOrPublisher="Fungal Diversity" pagination="101-117" refId="B16" refString="Divakar PK, Crespo A, Kraichak E, Leavitt SD, Singh G, Schmitt I, Lumbsch HT (2017) Using a temporal phylogenetic method to harmonize family- and genus-level classification in the largest clade of lichen-forming fungi. Fungal Diversity 84: 101117. https://doi.org/10.1007/s13225-017-0379-z" title="Using a temporal phylogenetic method to harmonize family- and genus-level classification in the largest clade of lichen-forming fungi." volume="84" year="2017">2017</bibRefCitation>
;
<bibRefCitation id="A454D7BB17BEFB25A47BB280454E8927" DOI="10.1017/S0024282911000570" author="Crespo" etAl="et al." firstAuthor="Crespo" journalOrPublisher="The Lichenologist" pagination="511-535" refId="B13" refString="Crespo A, Divakar PK, Hawksworth DL (2011) Generic concepts in parmelioid lichens, and the phylogenetic value of characters used in their circumscription. The Lichenologist 43: 511535. https://doi.org/10.1017/S0024282911000570" title="Generic concepts in parmelioid lichens, and the phylogenetic value of characters used in their circumscription." volume="43" year="2011">Crespo et al. 2011</bibRefCitation>
). As the species level taxonomy of
<taxonomicName id="D6DAC7D19327C2141944E6F60A1C4A06" class="Lecanoromycetes" family="Parmeliaceae" genus="Parmotrema" kingdom="Fungi" order="Lecanorales" phylum="Ascomycota" rank="genus">
<emphasis id="C3BFA4CD730A5F94EBE6B9D9935C6E58" italics="true">Parmotrema</emphasis>
</taxonomicName>
mainly relies on chemical and other characters not observable in the fossil inclusions, comparisons of fossils to extant species are often ineffectual. However, the genus currently includes several infrageneric groups that, at some point, have been acknowledged as separate genera based on morphology, anatomy, and chemistry but which were found to be nested within
<taxonomicName id="19552F0116E38C772FF2752A573EC335" class="Lecanoromycetes" family="Parmeliaceae" genus="Parmotrema" kingdom="Fungi" order="Lecanorales" phylum="Ascomycota" rank="genus">
<emphasis id="A27C7766CB64E1D194C73C59FE770525" italics="true">Parmotrema</emphasis>
</taxonomicName>
in molecular phylogenetic analyses (
<bibRefCitation id="D90F6AAC5518550F831AB0F738394766" DOI="10.2307/3243867" author="Elix" firstAuthor="Elix" refId="B19" refString="Elix JA (1993) Progress in the generic delimitation of Parmelia sensu lato lichens (Ascomycotina: Parmeliaceae) and a synoptic key to the Parmeliaceae. The Bryologist 96: 359-383. https://doi.org/10.2307/3243867" year="1993">Elix 1993</bibRefCitation>
;
<bibRefCitation id="798062E9CE42D7AFA71280E72AE3C2A4" DOI="10.1080/15572536.2006.11832848" author="Blanco" etAl="et al." firstAuthor="Blanco" journalOrPublisher="Mycologia" pagination="150-159" refId="B7" refString="Blanco O, Crespo A, Divakar PK, Elix JA, Lumbsch HT (2005) Molecular phylogeny of parmotremoid lichens (Ascomycota, Parmeliaceae). Mycologia 97: 150159. https://doi.org/10.1080/15572536.2006.11832848" title="Molecular phylogeny of parmotremoid lichens (Ascomycota, Parmeliaceae)." volume="97" year="2005">Blanco et al. 2005</bibRefCitation>
;
<bibRefCitation id="0EACBBB3ED1448F0781D6B3452B31F27" DOI="10.1017/S0024282904014586" author="Divakar" etAl="et al." firstAuthor="Divakar" journalOrPublisher="The Lichenologist" pagination="55-65" refId="B15" refString="Divakar PK, Blanco O, Hawksworth DL, Crespo A (2005) Molecular phylogenetic studies on the Parmotrema reticulatum (syn. Rimelia reticulata) complex, including the confirmation of P. pseudoreticulatum as a distinct species. The Lichenologist 37: 5565. https://doi.org/10.1017/S0024282904014586" title="Molecular phylogenetic studies on the Parmotrema reticulatum (syn. Rimelia reticulata) complex, including the confirmation of P. pseudoreticulatum as a distinct species." volume="37" year="2005">Divakar et al. 2005</bibRefCitation>
,
<bibRefCitation id="2F9A43990423E56C2D8E1859681C87A7" DOI="10.1007/s13225-017-0379-z" author="Divakar" firstAuthor="Divakar" journalOrPublisher="Fungal Diversity" pagination="101-117" refId="B16" refString="Divakar PK, Crespo A, Kraichak E, Leavitt SD, Singh G, Schmitt I, Lumbsch HT (2017) Using a temporal phylogenetic method to harmonize family- and genus-level classification in the largest clade of lichen-forming fungi. Fungal Diversity 84: 101117. https://doi.org/10.1007/s13225-017-0379-z" title="Using a temporal phylogenetic method to harmonize family- and genus-level classification in the largest clade of lichen-forming fungi." volume="84" year="2017">2017</bibRefCitation>
;
<bibRefCitation id="F37701CC179306034041EB70EA10D1E9" DOI="10.1017/S0024282911000570" author="Crespo" etAl="et al." firstAuthor="Crespo" journalOrPublisher="The Lichenologist" pagination="511-535" refId="B13" refString="Crespo A, Divakar PK, Hawksworth DL (2011) Generic concepts in parmelioid lichens, and the phylogenetic value of characters used in their circumscription. The Lichenologist 43: 511535. https://doi.org/10.1017/S0024282911000570" title="Generic concepts in parmelioid lichens, and the phylogenetic value of characters used in their circumscription." volume="43" year="2011">Crespo et al. 2011</bibRefCitation>
).
<taxonomicName id="7D7395708235BCF4AE223E80E08996F8" class="Lecanoromycetes" family="Parmeliaceae" genus="Parmotrema" kingdom="Fungi" order="Lecanorales" phylum="Ascomycota" rank="genus">
<emphasis id="E9F9A655456748C1111F2775D0036374" italics="true">Parmotrema</emphasis>
</taxonomicName>
specimen 1 with the smooth upper surface, dark lower surface, and very long, branched marginal cilia resembles the
<taxonomicName id="AA0016D8E2ADBA9E2D59E83A2C4AD01C" class="Lecanoromycetes" family="Parmeliaceae" genus="Parmotrema" kingdom="Fungi" order="Lecanorales" phylum="Ascomycota" rank="genus" sensu="stricto">
<emphasis id="B879A50D09E8367B0A41352A4DF2CEBB" italics="true">Parmotrema</emphasis>
</taxonomicName>
<taxonomicNameLabel id="082B64A92350C2B710F5F53AC7E09949" sensu="stricto">s. str.</taxonomicNameLabel>
group within the genus
<taxonomicName id="87B79DD41036FD4197EA70E8433EC6B3" class="Lecanoromycetes" family="Parmeliaceae" genus="Parmotrema" kingdom="Fungi" order="Lecanorales" phylum="Ascomycota" rank="genus">
<emphasis id="315F751188124C45030F87F031E3EA6F" italics="true">Parmotrema</emphasis>
</taxonomicName>
. The group is, for example, characterized by broad lobes, broad naked marginal zone on the lower surface, marginal cilia, and the lack of maculae on the upper surface (
<bibRefCitation id="C8759957834567139E28F53A532174D8" DOI="10.2307/3243867" author="Elix" firstAuthor="Elix" refId="B19" refString="Elix JA (1993) Progress in the generic delimitation of Parmelia sensu lato lichens (Ascomycotina: Parmeliaceae) and a synoptic key to the Parmeliaceae. The Bryologist 96: 359-383. https://doi.org/10.2307/3243867" year="1993">Elix 1993</bibRefCitation>
). The extant
<taxonomicName id="D5F478D48BA36118F7657D5C697A05CD" class="Lecanoromycetes" family="Parmeliaceae" genus="Parmotrema" kingdom="Fungi" order="Lecanorales" phylum="Ascomycota" rank="genus" sensu="stricto">
<emphasis id="99B4925F63752FE2BC1F676B6B7D92A4" italics="true">Parmotrema</emphasis>
</taxonomicName>
<taxonomicNameLabel id="6318D7D70F4B69DFFCD53AD93A7C6ABD" sensu="stricto">s. str.</taxonomicNameLabel>
group contains more than 250 species distributed especially in the tropical regions of the world (
<bibRefCitation id="0DF8C2EF572677157E795864B4EAFE9A" DOI="10.2307/3243867" author="Elix" firstAuthor="Elix" refId="B19" refString="Elix JA (1993) Progress in the generic delimitation of Parmelia sensu lato lichens (Ascomycotina: Parmeliaceae) and a synoptic key to the Parmeliaceae. The Bryologist 96: 359-383. https://doi.org/10.2307/3243867" year="1993">Elix 1993</bibRefCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>

View file

@ -0,0 +1,182 @@
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<mods:title id="3DA3109DCA9AB368C84968CAF3C5D5A4">Extending the fossil record of Miocene neotropical epiphyte communities</mods:title>
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<treatment id="CB108EEC9EB15CA981CFD92C76327C05" ID-arpha="CB108EEC-9EB1-5CA9-81CF-D92C76327C05" LSID="urn:lsid:plazi:treatment:CB108EEC9EB15CA981CFD92C76327C05" httpUri="http://treatment.plazi.org/id/CB108EEC9EB15CA981CFD92C76327C05" scope_class="Lecanoromycetes" scope_order="Lecanorales" scope_phylum="Ascomycota">
<subSubSection id="2063C3437EABCC2084E647422A73F6DE" type="nomenclature">
<paragraph id="12208E2060CCF6D06097AC4F83EF43C1">
<taxonomicName id="8BC26E7E3C999DB2313B9A0586B005AF" ID-CoL="6HY5" ID-ENA="172634" class="Lecanoromycetes" family="Parmeliaceae" genus="Parmotrema" kingdom="Fungi" order="Lecanorales" phylum="Ascomycota" rank="species" species="undetermined-2">
<emphasis id="D2AC6D474296B5B2B717DA89A2586057" italics="true">Parmotrema</emphasis>
specimen 2
</taxonomicName>
</paragraph>
</subSubSection>
<subSubSection id="SECID0ELJBI" type="Specimen">
<paragraph id="E7BF869C1B1F5AB58D91DA129FF6EB5E">
<heading id="EDDA72A6F9A7AD6D37BC5D46BE3E6D42" reason="title">Specimen.</heading>
</paragraph>
<paragraph id="409A414D4F6D7A413250566138050248">
Stuttgart State Museum of Natural History,
<collectingCountry id="253ABCC2F1CF4A26B6141D8378A2060B" name="Germany">Germany</collectingCountry>
(
<named-content id="5385CAB155ED529FE36A5191B3B32C75" content-type="dwc:institutional_code" xlink_href="http://grbio.org/institution/staatliches-museum-fuer-naturkund-stuttgart" xlink_title="Stuttgart State Museum of Natural History, Germany">
<collectionCode id="B73E0AFC7A433CEECA24C4D7EAE6DA22" country="Germany" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34840" lsid="urn:lsid:biocol.org:col:34840" name="Staatliches Museum fuer Naturkund Stuttgart" type="">SMNS</collectionCode>
</named-content>
),
<specimenCode id="9694F6BA97859AE5620846E516B29EA0">
<named-content id="B845D014F03B05BEB2B98BDD95F0D54F" content-type="dwc:institutional_code" xlink_href="http://grbio.org/institution/staatliches-museum-fuer-naturkund-stuttgart" xlink_title="Stuttgart State Museum of Natural History, Germany">SMNS</named-content>
-DO-4929-M
</specimenCode>
.
</paragraph>
</subSubSection>
<subSubSection id="SECID0E2JBI" type="Age and stratigraphic level">
<paragraph id="C38159F9AF690B21190668B9E6A5F743">
<heading id="E87D8D1518BB8BEA3E9742D742560A89" reason="title">Age and stratigraphic level.</heading>
</paragraph>
<paragraph id="D05ECCF641141C81B3C5BCDB1F3BE9D8">
1520 Ma, Langhian Burdigalian (early to middle Miocene), La Toca Formation,
<collectingCountry id="FE9A80FBD82E54D056B4BF69EC91D09A" name="Dominican Republic">Dominican Republic</collectingCountry>
.
</paragraph>
</subSubSection>
<subSubSection id="SECID0EKKBI" type="description">
<paragraph id="6C5EC613ED9C5031E1CC31F75EB98DAE">
<heading id="45BAA23AF5940C48D76EDF3AEAA6B1A3" reason="title">Description.</heading>
</paragraph>
<paragraph id="66B7178A644CABDDEB561EF90A673B18">
Three lichen fragments situated very closely together, most probably originating from the same thallus (Fig.
<figureCitation id="EB02E8FC191F5A20B1D0FBF5AB98A593" captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. Parmotrema fossils from Dominican amber. A-C. Parmotrema specimen 1 (SMNS - DO- 4928 - M); A. Parmotrema inclusion with a bryophyte (leafy liverwort, Porellales); B. Lobes seen from above showing the truncate lobe apices, dark lobe margins, and long marginal cilia. The lighter spots visible on the upper surface of the thallus are caused by slight depressions and their brighter color derives from air-filled spaces between the thallus surface and amber. These depressions seem to correspond to the attachment points of the rhizines on the lower surface; C. Close-up of the slightly uneven upper surface and finger-like isidia. D-E. Parmotrema specimen 2 (SMNS - DO- 4929 - M); D. Close-up of the upper surface with abundant pycnidia; E. General view of the inclusion, showing the very long, branching marginal cilia." figureDoi="10.3897/fr.28.137758.figure7" httpUri="https://binary.pensoft.net/fig/1264819">7 E</figureCitation>
); whole inclusion approximately 5.0 ×
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in diameter.
<emphasis id="366111005B61995C3B707647DEB59A29" bold="true" italics="true">Thallus</emphasis>
foliose, lobate.
<emphasis id="668CB063937D0555110E6B86F667A822" bold="true" italics="true">Lobes</emphasis>
flat, broad to linear, 0.41.0 mm wide, lobe apices truncate to rounded (Fig.
<figureCitation id="ABC897895AF2292C99F3A444C613E097" captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. Parmotrema fossils from Dominican amber. A-C. Parmotrema specimen 1 (SMNS - DO- 4928 - M); A. Parmotrema inclusion with a bryophyte (leafy liverwort, Porellales); B. Lobes seen from above showing the truncate lobe apices, dark lobe margins, and long marginal cilia. The lighter spots visible on the upper surface of the thallus are caused by slight depressions and their brighter color derives from air-filled spaces between the thallus surface and amber. These depressions seem to correspond to the attachment points of the rhizines on the lower surface; C. Close-up of the slightly uneven upper surface and finger-like isidia. D-E. Parmotrema specimen 2 (SMNS - DO- 4929 - M); D. Close-up of the upper surface with abundant pycnidia; E. General view of the inclusion, showing the very long, branching marginal cilia." figureDoi="10.3897/fr.28.137758.figure7" httpUri="https://binary.pensoft.net/fig/1264819">7 E</figureCitation>
).
<emphasis id="0471E3B7FDD3541C9AF170DDEE45790B" bold="true" italics="true">Upper surface</emphasis>
smooth and brown, with dark margins (Fig.
<figureCitation id="46FEE78A5DAFB5834C959EBD6746D891" captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. Parmotrema fossils from Dominican amber. A-C. Parmotrema specimen 1 (SMNS - DO- 4928 - M); A. Parmotrema inclusion with a bryophyte (leafy liverwort, Porellales); B. Lobes seen from above showing the truncate lobe apices, dark lobe margins, and long marginal cilia. The lighter spots visible on the upper surface of the thallus are caused by slight depressions and their brighter color derives from air-filled spaces between the thallus surface and amber. These depressions seem to correspond to the attachment points of the rhizines on the lower surface; C. Close-up of the slightly uneven upper surface and finger-like isidia. D-E. Parmotrema specimen 2 (SMNS - DO- 4929 - M); D. Close-up of the upper surface with abundant pycnidia; E. General view of the inclusion, showing the very long, branching marginal cilia." figureDoi="10.3897/fr.28.137758.figure7" httpUri="https://binary.pensoft.net/fig/1264819">7 D</figureCitation>
).
<emphasis id="F41A7A0BBDB86AB006ABD2ACE474CDE8" bold="true" italics="true">Marginal cilia</emphasis>
dark, branched, and long (up to
<quantity id="7D8ACAE34B00888F85A085C1D34F45DF" metricMagnitude="-4" metricUnit="m" metricValue="8.0" unit="mm" value="0.8">0.8 mm</quantity>
) (Fig.
<figureCitation id="235EB735741F1CB64E819FE137933E89" captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. Parmotrema fossils from Dominican amber. A-C. Parmotrema specimen 1 (SMNS - DO- 4928 - M); A. Parmotrema inclusion with a bryophyte (leafy liverwort, Porellales); B. Lobes seen from above showing the truncate lobe apices, dark lobe margins, and long marginal cilia. The lighter spots visible on the upper surface of the thallus are caused by slight depressions and their brighter color derives from air-filled spaces between the thallus surface and amber. These depressions seem to correspond to the attachment points of the rhizines on the lower surface; C. Close-up of the slightly uneven upper surface and finger-like isidia. D-E. Parmotrema specimen 2 (SMNS - DO- 4929 - M); D. Close-up of the upper surface with abundant pycnidia; E. General view of the inclusion, showing the very long, branching marginal cilia." figureDoi="10.3897/fr.28.137758.figure7" httpUri="https://binary.pensoft.net/fig/1264819">7 E</figureCitation>
).
<emphasis id="AD61FC2063D900C04569FBF3462972BB" bold="true" italics="true">Medulla</emphasis>
not visible.
<emphasis id="A6463E5169303035A7577249CC6D0CCE" bold="true" italics="true">Lower cortex</emphasis>
dark.
<emphasis id="8D239E2A7A907D764A56C39BAE056CE0" bold="true" italics="true">Rhizines</emphasis>
dark.
<emphasis id="777C69D3B6B43EEC52326B14BB6AE0F8" bold="true" italics="true">Pycnidia</emphasis>
abundant on the upper surface (Fig.
<figureCitation id="9998B412E34455A8BAEA2EDD7F43B2BA" captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. Parmotrema fossils from Dominican amber. A-C. Parmotrema specimen 1 (SMNS - DO- 4928 - M); A. Parmotrema inclusion with a bryophyte (leafy liverwort, Porellales); B. Lobes seen from above showing the truncate lobe apices, dark lobe margins, and long marginal cilia. The lighter spots visible on the upper surface of the thallus are caused by slight depressions and their brighter color derives from air-filled spaces between the thallus surface and amber. These depressions seem to correspond to the attachment points of the rhizines on the lower surface; C. Close-up of the slightly uneven upper surface and finger-like isidia. D-E. Parmotrema specimen 2 (SMNS - DO- 4929 - M); D. Close-up of the upper surface with abundant pycnidia; E. General view of the inclusion, showing the very long, branching marginal cilia." figureDoi="10.3897/fr.28.137758.figure7" httpUri="https://binary.pensoft.net/fig/1264819">7 D</figureCitation>
). Apothecia, isidia, or soredia not present.
</paragraph>
</subSubSection>
<subSubSection id="SECID0ELMBI" type="discussion">
<paragraph id="9C613712569F312298629BD621EC5980">
<heading id="F735A75DA28EBCCC55949707DC6A5425" reason="title">Discussion.</heading>
</paragraph>
<paragraph id="9F9295D81D7B1CABAB5EF28A5CA06D46">
The general habit, very long, branched marginal cilia, and dark lower surface identify the specimen as
<taxonomicName id="146F4B9C242779B3FBE4E973685F0D45" class="Lecanoromycetes" family="Parmeliaceae" genus="Parmotrema" kingdom="Fungi" order="Lecanorales" phylum="Ascomycota" rank="genus">
<emphasis id="26E717F239936B6027CBA49BD96CE986" italics="true">Parmotrema</emphasis>
</taxonomicName>
. Of the infrageneric groups, the dark lower surface without rhizines reaching the margins could represent, for example, the
<taxonomicName id="C3C4E9A6372248766DAB27F169C5644C" class="Lecanoromycetes" family="Parmeliaceae" genus="Parmotrema" kingdom="Fungi" order="Lecanorales" phylum="Ascomycota" rank="genus" sensu="stricto">
<emphasis id="042757308E4235BB095CA6FC5050D8E5" italics="true">Parmotrema</emphasis>
</taxonomicName>
<taxonomicNameLabel id="B218B4819DAE01B0BEF2EFEBF285A051" sensu="stricto">s. str.</taxonomicNameLabel>
group.
</paragraph>
</subSubSection>
</treatment>
</document>

View file

@ -0,0 +1,241 @@
<document id="E0BC8D34A5EB4533EAD59772D715FA56" ID-DOI="10.3897/fr.28.137758" ID-ZooBank="A10C1262-F859-4004-A83C-C8F032C1156A" ID-publisher-id="137758" URI-arpha="66B09103-4195-59B0-BBF5-6B2263A772A9" URI-zoobank="http://zoobank.org/A10C1262-F859-4004-A83C-C8F032C1156A" XM.bibliography_approvedBy="admin" XM.materialsCitations_approvedBy="admin" XM.taxonomicNames_approvedBy="admin" article-type="research-article" checkinTime="1740755594576" checkinUser="pensoft" docAuthor="Feldberg, Kathrin, Kaasalainen, Ulla, Mamontov, Yuriy S., Gradstein, S. Robbert, Schäfer-Verwimp, Alfons, Divakar, Pradeep K. &amp; Schmidt, Alexander R." docDate="2025" docId="FB66CEF003875E48827155D844A39F21" docLanguage="en" docName="Fossil Record 28(1): 79-102" docOrigin="Fossil Record 28 (1)" docSource="https://fr.pensoft.net/article/137758/download/xml/" docStyle="DocumentStyle:PensoftTaxPub.0000.journal_article.generic" docStyleName="PensoftTaxPub.0000.journal_article.generic" docTitle="Parmotrema undetermined-3" docType="treatment" docVersion="2" dtd-version="3.0" lastPageNumber="102" masterDocId="66B09103419559B0BBF56B2263A772A9" masterDocTitle="Extending the fossil record of Miocene neotropical epiphyte communities" masterLastPageNumber="102" masterPageNumber="79" pageNumber="79" updateTime="1740768570675" updateUser="admin">
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<mods:titleInfo id="7ED046CAC7A5CA03A84C32650BEECD1D">
<mods:title id="44247424B246C81DFD9952EBD8DD50AB">Extending the fossil record of Miocene neotropical epiphyte communities</mods:title>
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<mods:name id="A6713D8DAF1B45070F132F5F97D44A51" type="personal">
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<mods:namePart id="F72D605667D47E12AFC4065098390953">Feldberg, Kathrin</mods:namePart>
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<mods:affiliation id="30BB4C21FFBA4DE844260C091550391C">Department of Geobiology, University of Göttingen, Goldschmidtstraße 3, 37077 Göttingen, Germany</mods:affiliation>
</mods:name>
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<mods:namePart id="76445A9D01211BC1D75DF895A2CBC797">Kaasalainen, Ulla</mods:namePart>
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<mods:affiliation id="49AECF1BFFC09C4369B7DAB2C6597AE5">Finnish Museum of Natural History, University of Helsinki, P. O. Box 7, 00014 Helsinki, Finland</mods:affiliation>
</mods:name>
<mods:name id="E1C566C0FFBC3AC7895FBE78957470C6" type="personal">
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<mods:namePart id="72948F3ED5737D4AD3C837D61ADAE497">Mamontov, Yuriy S.</mods:namePart>
<mods:nameIdentifier id="7A9512487022ADD5D7EB9430724A8683" type="ORCID">0000-0003-3851-0738</mods:nameIdentifier>
<mods:affiliation id="79E20304DD5DE57474299A9D1F35EE94">Tsitsin Main Botanical Garden, Russian Academy of Sciences, Moscow, 127276 Russia</mods:affiliation>
</mods:name>
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<mods:role id="350432CE4CB09E47115F300EA409431A">
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<mods:namePart id="24158D738B68C41B01B6A55108EDAD93">Gradstein, S. Robbert</mods:namePart>
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<mods:affiliation id="706F96C4D9D5F4D15BAF01440A3E88C3">Meise Botanic Garden, 1860 Meise, Belgium</mods:affiliation>
</mods:name>
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</mods:role>
<mods:namePart id="0A243F3C8575BF86632AB645A3603827">Schäfer-Verwimp, Alfons</mods:namePart>
<mods:nameIdentifier id="211EEC815DC13ADA3C3D10198E09ABC4" type="ORCID">0000-0002-2720-6055</mods:nameIdentifier>
<mods:affiliation id="EF2EFA3F033D4EF7B77409658696AF6B">Mittlere Letten 11, 88634 Herdwangen-Schönach, Germany</mods:affiliation>
</mods:name>
<mods:name id="BCFB859A4097B3BE53E8745C5ADFCB7B" type="personal">
<mods:role id="E8A2055734FEFF096DEA895163A332BA">
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</mods:role>
<mods:namePart id="6E25BF9DE64C41EDA9BA4927FF64440E">Divakar, Pradeep K.</mods:namePart>
<mods:nameIdentifier id="2341DBB948ED709D65EBCFFD2F9E8B1D" type="ORCID">0000-0002-0300-0124</mods:nameIdentifier>
<mods:affiliation id="FFE8022DEC5AF3F7972A53A45751DC64">Departamento de Biología Vegetal II, Facultad de Farmacia, Universidad Complutense de Madrid, 28040 Madrid, Spain</mods:affiliation>
</mods:name>
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</mods:role>
<mods:namePart id="EECB36B4888F21E7AFB689FD2586141D">Schmidt, Alexander R.</mods:namePart>
<mods:nameIdentifier id="E33D0236672A08E5695D439EE854F3E1" type="ORCID">0000-0001-5426-4667</mods:nameIdentifier>
<mods:affiliation id="A32CAED1B0B5B2794C7A8332FF266C1D">Department of Geobiology, University of Göttingen, Goldschmidtstraße 3, 37077 Göttingen, Germany</mods:affiliation>
</mods:name>
<mods:typeOfResource id="F1B6A7A6C5298B8B0E047AEAE167B600">text</mods:typeOfResource>
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<mods:titleInfo id="6CADECA70E67B6C344EB06494C951411">
<mods:title id="2706D29AB4C7565BA32A6CBB04AE9D76">Fossil Record</mods:title>
</mods:titleInfo>
<mods:part id="3227465017932738987243B45008A4DD">
<mods:date id="CBCAEC73D0023ACC2753DD94439FB78B">2025</mods:date>
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<mods:number id="C54087BEB8416451B8A81BA678AF2FA3">2025-02-27</mods:number>
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<mods:number id="C3963425A095DC8A65CE84C4FEB2ABEB">28</mods:number>
</mods:detail>
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<mods:number id="B9EA64BEABC060F26829B474B2D4D840">1</mods:number>
</mods:detail>
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<mods:start id="D801BB4B64ECCDC97A487FF3AF2903E6">79</mods:start>
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</mods:extent>
</mods:part>
</mods:relatedItem>
<mods:classification id="9A3AE67302355710E4D540760961DA91">journal article</mods:classification>
<mods:identifier id="AF50D9ABC69D9054DB19E20AAD9909CC" type="DOI">10.3897/fr.28.137758</mods:identifier>
<mods:identifier id="08D2B7E5B2233C9AEB0C608FE6A1B5CA" type="ZooBank">A10C1262-F859-4004-A83C-C8F032C1156A</mods:identifier>
</mods:mods>
<treatment id="FB66CEF003875E48827155D844A39F21" ID-arpha="FB66CEF0-0387-5E48-8271-55D844A39F21" LSID="urn:lsid:plazi:treatment:FB66CEF003875E48827155D844A39F21" httpUri="http://treatment.plazi.org/id/FB66CEF003875E48827155D844A39F21" scope_class="Lecanoromycetes" scope_order="Lecanorales" scope_phylum="Ascomycota">
<subSubSection id="6CE2BD173EC0F3B07ED9BDDAD96D1AB2" type="nomenclature">
<paragraph id="36FBA90676CDDE88C7FB57BA6D66C7F0">
<taxonomicName id="3337FA7D44649AEAB55340BF8AA32C97" ID-CoL="6HY5" ID-ENA="172634" class="Lecanoromycetes" family="Parmeliaceae" genus="Parmotrema" kingdom="Fungi" order="Lecanorales" phylum="Ascomycota" rank="species" species="undetermined-3">
<emphasis id="2A076C33AA8C2E5198CDF256BC0C4DBE" italics="true">Parmotrema</emphasis>
specimen 3
</taxonomicName>
</paragraph>
</subSubSection>
<subSubSection id="SECID0E6NBI" type="Specimen">
<paragraph id="CC44A48B4E23ED60AF040E0A95E08B5B">
<heading id="1DE487BE9CD7F676F499E4D67DDDFF19" reason="title">Specimen.</heading>
</paragraph>
<paragraph id="62E059864A734678C2C803FAAABD57FD">
American Museum of Natural History, New
<collectingRegion id="8011CBEC26A4A314661371CEBE40EF01" country="United Kingdom" name="York">York</collectingRegion>
,
<collectionCode id="3986DA92C0063018C5D7E73D6F303A39" country="USA" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34925" lsid="urn:lsid:biocol.org:col:34925" name="American Museum of Natural History" type="">AMNH</collectionCode>
<specimenCode id="C8F1F55DEFF58102BEE58B38990DD575">DR-14-294</specimenCode>
.
</paragraph>
</subSubSection>
<subSubSection id="SECID0EEOBI" type="Age and stratigraphic level">
<paragraph id="588BA44CCB2917A01CA9D3F092009235">
<heading id="BA348586C751017631A13FF5D8E8B5D3" reason="title">Age and stratigraphic level.</heading>
</paragraph>
<paragraph id="0A9951E5A2B945E3546747AF183F90EA">
1520 Ma, Langhian Burdigalian (early to middle Miocene), La Toca Formation,
<collectingCountry id="59651E018504A970F626E1FC612041D7" name="Dominican Republic">Dominican Republic</collectingCountry>
.
</paragraph>
</subSubSection>
<subSubSection id="SECID0ETOBI" type="description">
<paragraph id="1C3E9D879592EF8C219221451C79590B">
<heading id="B43561A5884B0C523D76725C015FB415" reason="title">Description.</heading>
</paragraph>
<paragraph id="FCA54DBECD5642EB44E5FE0EE520767A">
Lichen fragment approximately 1.7 ×
<quantity id="923F7F99EF01899DB45C74CBB51AEA19" metricMagnitude="-3" metricUnit="m" metricValue="1.3" unit="mm" value="1.3">1.3 mm</quantity>
in diameter.
<emphasis id="9987BE1D5AB4A3D0359DA5293953F1E5" bold="true" italics="true">Thallus</emphasis>
foliose, lobate.
<emphasis id="882FCEA7252DA0429D7D30082C41E251" bold="true" italics="true">Lobes</emphasis>
flat, linear, approximately
<quantity id="5EE429C2A0DFD8BD04DA6A3A36B0E280" metricMagnitude="-4" metricUnit="m" metricValue="5.0" unit="mm" value="0.5">0.5 mm</quantity>
wide and 40 µm thick, lobe apices truncate to rounded.
<emphasis id="DD61CF2F3AFBD4CEFADC7F7FF74D8311" bold="true" italics="true">Upper surface</emphasis>
slightly uneven with effigurate maculae, with prominent dark margins (Fig.
<figureCitation id="D278A9ACDF830E384C94917375AB0D0B" captionStart="Figure 8" captionStartId="F8" captionText="Figure 8. Parmotrema fossils from Dominican and Mexican ambers. A-C. Parmotrema specimen 3 (AMNH DR- 14-294) from Dominican amber; A. Maculate upper surface and long, branching marginal cilia; B. Lower surface with rhizines; C. Dark lower cortex and a cross-section of the lobe revealing the medulla; D, E. Parmotrema specimen 4 (GZG. BST. 22085) from Mexican amber; D. Smooth upper surface and long, dark marginal cilia. The dark, robust rhizines are also visible from the upper side through the semitransparent thallus; E. Branching rhizines on the lower surface." figureDoi="10.3897/fr.28.137758.figure8" httpUri="https://binary.pensoft.net/fig/1264820">8 A</figureCitation>
).
<emphasis id="1CE9E40D9A648BD99691007E8B03F8B1" bold="true" italics="true">Marginal cilia</emphasis>
long (up to
<quantity id="5F53494E37F9B2D16FBA29F0DC57CC3A" metricMagnitude="-4" metricUnit="m" metricValue="9.0" unit="mm" value="0.9">0.9 mm</quantity>
), branching, tapering (Fig.
<figureCitation id="15CED53D69F65DE0CC9B6D2866F2B490" captionStart="Figure 8" captionStartId="F8" captionText="Figure 8. Parmotrema fossils from Dominican and Mexican ambers. A-C. Parmotrema specimen 3 (AMNH DR- 14-294) from Dominican amber; A. Maculate upper surface and long, branching marginal cilia; B. Lower surface with rhizines; C. Dark lower cortex and a cross-section of the lobe revealing the medulla; D, E. Parmotrema specimen 4 (GZG. BST. 22085) from Mexican amber; D. Smooth upper surface and long, dark marginal cilia. The dark, robust rhizines are also visible from the upper side through the semitransparent thallus; E. Branching rhizines on the lower surface." figureDoi="10.3897/fr.28.137758.figure8" httpUri="https://binary.pensoft.net/fig/1264820">8 A</figureCitation>
).
<emphasis id="D19E6C5E01AF2FE6696ACA0D5FF3A388" bold="true" italics="true">Medulla</emphasis>
pale (Fig.
<figureCitation id="CA4B74B85C9EAED68600A54BF644CDC2" captionStart="Figure 8" captionStartId="F8" captionText="Figure 8. Parmotrema fossils from Dominican and Mexican ambers. A-C. Parmotrema specimen 3 (AMNH DR- 14-294) from Dominican amber; A. Maculate upper surface and long, branching marginal cilia; B. Lower surface with rhizines; C. Dark lower cortex and a cross-section of the lobe revealing the medulla; D, E. Parmotrema specimen 4 (GZG. BST. 22085) from Mexican amber; D. Smooth upper surface and long, dark marginal cilia. The dark, robust rhizines are also visible from the upper side through the semitransparent thallus; E. Branching rhizines on the lower surface." figureDoi="10.3897/fr.28.137758.figure8" httpUri="https://binary.pensoft.net/fig/1264820">8 C</figureCitation>
).
<emphasis id="FCAF2C1E5508B09F15CACAFDC5D17EF9" bold="true" italics="true">Lower cortex</emphasis>
dark (Fig.
<figureCitation id="D29C433E48D2A8FA444B67FCDB2B135E" captionStart="Figure 8" captionStartId="F8" captionText="Figure 8. Parmotrema fossils from Dominican and Mexican ambers. A-C. Parmotrema specimen 3 (AMNH DR- 14-294) from Dominican amber; A. Maculate upper surface and long, branching marginal cilia; B. Lower surface with rhizines; C. Dark lower cortex and a cross-section of the lobe revealing the medulla; D, E. Parmotrema specimen 4 (GZG. BST. 22085) from Mexican amber; D. Smooth upper surface and long, dark marginal cilia. The dark, robust rhizines are also visible from the upper side through the semitransparent thallus; E. Branching rhizines on the lower surface." figureDoi="10.3897/fr.28.137758.figure8" httpUri="https://binary.pensoft.net/fig/1264820">8 B, C</figureCitation>
).
<emphasis id="5E2EC2B662E64E6A5B9A57F2AD677850" bold="true" italics="true">Rhizines</emphasis>
dark, mainly simple (Fig.
<figureCitation id="C011DFD65C08254EA020792DE313D7E7" captionStart="Figure 8" captionStartId="F8" captionText="Figure 8. Parmotrema fossils from Dominican and Mexican ambers. A-C. Parmotrema specimen 3 (AMNH DR- 14-294) from Dominican amber; A. Maculate upper surface and long, branching marginal cilia; B. Lower surface with rhizines; C. Dark lower cortex and a cross-section of the lobe revealing the medulla; D, E. Parmotrema specimen 4 (GZG. BST. 22085) from Mexican amber; D. Smooth upper surface and long, dark marginal cilia. The dark, robust rhizines are also visible from the upper side through the semitransparent thallus; E. Branching rhizines on the lower surface." figureDoi="10.3897/fr.28.137758.figure8" httpUri="https://binary.pensoft.net/fig/1264820">8 B</figureCitation>
). Apothecia, isidia, soredia, or pycnidia not present.
</paragraph>
<caption id="5DE70225E8EDAEB02E87175D98A99B3B" ID-DOI="10.3897/fr.28.137758.figure8" ID-arpha="9B1C827C-A02F-5366-862A-37CB433992CA" httpUri="https://binary.pensoft.net/fig/1264820" startId="F8">
<paragraph id="27F08A571CC14996045A56DD76EE2A23">
<label id="651EBD9E90D9757683452622AEE74A9C">Figure 8.</label>
</paragraph>
<paragraph id="FD6788BE3510EE6F6DABFF726DF55901">
<taxonomicName id="E747FA53B45C2E757E79F8DFD24AE2BD" class="Lecanoromycetes" family="Parmeliaceae" genus="Parmotrema" kingdom="Fungi" order="Lecanorales" phylum="Ascomycota" rank="genus">
<emphasis id="FE7F740667D1DEFE5C5DB9D647E783E9" italics="true">Parmotrema</emphasis>
</taxonomicName>
fossils from Dominican and Mexican ambers.
<emphasis id="D933A36F35A94FA4FD182C652C22C834" bold="true">A-C.</emphasis>
<taxonomicName id="58254BF44BE01A64A9F02E4E3B250D94" class="Lecanoromycetes" family="Parmeliaceae" genus="Parmotrema" kingdom="Fungi" order="Lecanorales" phylum="Ascomycota" rank="genus">
<emphasis id="2095FC471E4835E0EEF43715F3EDB540" italics="true">Parmotrema</emphasis>
</taxonomicName>
specimen 3 (AMNH
<specimenCode id="0C952F513C7BF6F9BB7B11B689D4CE44">DR-14-294</specimenCode>
) from Dominican amber;
<emphasis id="93E20959DEAA7C1090DBE6A32706B7BC" bold="true">A.</emphasis>
Maculate upper surface and long, branching marginal cilia;
<emphasis id="99C58FFE17D3773D474B92E82F539B1E" bold="true">B.</emphasis>
Lower surface with rhizines;
<emphasis id="12A622637CEA27EF1DDDA89CB6CCD0D7" bold="true">C.</emphasis>
Dark lower cortex and a cross-section of the lobe revealing the medulla;
<emphasis id="0504E8BD9C1224D27446804B29E06900" bold="true">D, E.</emphasis>
<taxonomicName id="2CB37A582D6E0AB5E40812ADF91CE077" class="Lecanoromycetes" family="Parmeliaceae" genus="Parmotrema" kingdom="Fungi" order="Lecanorales" phylum="Ascomycota" rank="genus">
<emphasis id="D505B8EEA3CF0D3FD3066D5561BA3EA2" italics="true">Parmotrema</emphasis>
</taxonomicName>
specimen 4 (
<specimenCode id="98B0DF2B196229A88DDFFB59376966E3">
<abbrev content-type="institution" id="ABBRID0EDSBI" xlink_title="Geoscientific Collection of the University of Göttingen">GZG</abbrev>
.BST.22085
</specimenCode>
) from Mexican amber;
<emphasis id="766200839A16AF6D05994156296ADACE" bold="true">D.</emphasis>
Smooth upper surface and long, dark marginal cilia. The dark, robust rhizines are also visible from the upper side through the semitransparent thallus;
<emphasis id="7FE73D7B171B2E79E3F903FC32557A08" bold="true">E.</emphasis>
Branching rhizines on the lower surface.
</paragraph>
</caption>
</subSubSection>
<subSubSection id="SECID0EXSBI" type="discussion">
<paragraph id="945E0BEF356FD67B613A0B8DA2296737">
<heading id="0C96E06DD47FB42200F371116C955CF1" reason="title">Discussion.</heading>
</paragraph>
<paragraph id="4BE5B5FB39FA496CF698F0E8389890BB">
The general habit and long, branching and tapering marginal cilia identify the specimen as
<taxonomicName id="6F8050647C934FDD072592831C22DC62" class="Lecanoromycetes" family="Parmeliaceae" genus="Parmotrema" kingdom="Fungi" order="Lecanorales" phylum="Ascomycota" rank="genus">
<emphasis id="FC0FD24C59CD8712491766D2E7430898" italics="true">Parmotrema</emphasis>
</taxonomicName>
. The effigurate maculae on the upper surface and tapered marginal cilia of
<taxonomicName id="65413A93794D4A46D601DBA6F5D1E78A" class="Lecanoromycetes" family="Parmeliaceae" genus="Parmotrema" kingdom="Fungi" order="Lecanorales" phylum="Ascomycota" rank="genus">
<emphasis id="55928DFAECD70C47795B84CF10CA4F88" italics="true">Parmotrema</emphasis>
</taxonomicName>
specimen 3 resemble the former genus
<taxonomicName id="02DFE0AC77467FB955B106ED5E76EEAD" authority="(Elix 1993)" baseAuthorityName="Elix" baseAuthorityYear="1993" class="Lecanoromycetes" family="Parmeliaceae" genus="Canomaculina" kingdom="Fungi" order="Lecanorales" phylum="Ascomycota" rank="genus">
<emphasis id="752EC6C1EB0DAC10B17EA0C88157C828" italics="true">Canomaculina</emphasis>
(
<bibRefCitation id="37335CC6C196D8A4CA03B66DA43EFA75" DOI="10.2307/3243867" author="Elix" firstAuthor="Elix" refId="B19" refString="Elix JA (1993) Progress in the generic delimitation of Parmelia sensu lato lichens (Ascomycotina: Parmeliaceae) and a synoptic key to the Parmeliaceae. The Bryologist 96: 359-383. https://doi.org/10.2307/3243867" year="1993">Elix 1993</bibRefCitation>
)
</taxonomicName>
which has been synonymized with
<taxonomicName id="CCBB7F475F4B8857E4A986C6AB811624" authority="(Blanco et al. 2005)" baseAuthorityName="Blanco" baseAuthorityYear="2005" class="Lecanoromycetes" family="Parmeliaceae" genus="Parmotrema" kingdom="Fungi" order="Lecanorales" phylum="Ascomycota" rank="genus">
<emphasis id="8A6EC3454878D5CA9ED4116F2E31666C" italics="true">Parmotrema</emphasis>
(
<bibRefCitation id="5450B7510DB32437A11B547F14C95202" DOI="10.1080/15572536.2006.11832848" author="Blanco" etAl="et al." firstAuthor="Blanco" journalOrPublisher="Mycologia" pagination="150-159" refId="B7" refString="Blanco O, Crespo A, Divakar PK, Elix JA, Lumbsch HT (2005) Molecular phylogeny of parmotremoid lichens (Ascomycota, Parmeliaceae). Mycologia 97: 150159. https://doi.org/10.1080/15572536.2006.11832848" title="Molecular phylogeny of parmotremoid lichens (Ascomycota, Parmeliaceae)." volume="97" year="2005">Blanco et al. 2005</bibRefCitation>
)
</taxonomicName>
. The
<taxonomicName id="EB27051EC91FA0197C4B4729C8C43F94" baseAuthorityName="Elix" baseAuthorityYear="1993" class="Lecanoromycetes" family="Parmeliaceae" genus="Canomaculina" kingdom="Fungi" order="Lecanorales" phylum="Ascomycota" rank="genus">
<emphasis id="1F1452994E0EAC993235FB099161831A" italics="true">Canomaculina</emphasis>
</taxonomicName>
group is characterized by tapered cilia, effigurate-maculate upper surface of the lobes, and simple and furcate to branched rhizines (
<bibRefCitation id="F21B02D89EDD50AC3ED273E6401D2A43" DOI="10.2307/3243867" author="Elix" firstAuthor="Elix" refId="B19" refString="Elix JA (1993) Progress in the generic delimitation of Parmelia sensu lato lichens (Ascomycotina: Parmeliaceae) and a synoptic key to the Parmeliaceae. The Bryologist 96: 359-383. https://doi.org/10.2307/3243867" year="1993">Elix 1993</bibRefCitation>
). The
<taxonomicName id="D8794A4FFE0C328EF0137F211EF45492" baseAuthorityName="Elix" baseAuthorityYear="1993" class="Lecanoromycetes" family="Parmeliaceae" genus="Canomaculina" kingdom="Fungi" order="Lecanorales" phylum="Ascomycota" rank="genus">
<emphasis id="01EB6BD99A0B33B921DF4BE92FB67763" italics="true">Canomaculina</emphasis>
</taxonomicName>
group contains more than ten extant species and has a center of distribution in South America (
<bibRefCitation id="D632B718D43381012C91884197FCDFE8" DOI="10.2307/3243867" author="Elix" firstAuthor="Elix" refId="B19" refString="Elix JA (1993) Progress in the generic delimitation of Parmelia sensu lato lichens (Ascomycotina: Parmeliaceae) and a synoptic key to the Parmeliaceae. The Bryologist 96: 359-383. https://doi.org/10.2307/3243867" year="1993">Elix 1993</bibRefCitation>
;
<bibRefCitation id="A51F2EDAADBD28FBBE10E1C38529E2AA" DOI="10.1080/15572536.2006.11832848" author="Blanco" etAl="et al." firstAuthor="Blanco" journalOrPublisher="Mycologia" pagination="150-159" refId="B7" refString="Blanco O, Crespo A, Divakar PK, Elix JA, Lumbsch HT (2005) Molecular phylogeny of parmotremoid lichens (Ascomycota, Parmeliaceae). Mycologia 97: 150159. https://doi.org/10.1080/15572536.2006.11832848" title="Molecular phylogeny of parmotremoid lichens (Ascomycota, Parmeliaceae)." volume="97" year="2005">Blanco et al. 2005</bibRefCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>