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<document id="BA35FF47AE192A7F2C8E59CA27D81DF9" ID-DOI="10.11646/phytotaxa.652.4.4" ID-ISSN="1179-3163" ID-Zenodo-Dep="13216837" IM.bibliography_approvedBy="carolina" IM.illustrations_approvedBy="felipe" IM.materialsCitations_approvedBy="carolina" IM.metadata_approvedBy="carolina" IM.taxonomicNames_approvedBy="carolina" IM.treatments_approvedBy="felipe" checkinTime="1722603772051" checkinUser="felipe" docAuthor="Kociolek, J. Patrick, Greenwood, Megan, Rogers, Dillan, Hamsher, Sarah E., Miller, Scott, Li, Jingchun, Chang, Aimee Caye G. &amp; Taylor, Jonathan" docDate="2024" docId="EA548784FFA1B073FF0C1AFBFDDD1D47" docLanguage="en" docName="phytotaxa.652.4.4.pdf" docOrigin="Phytotaxa 652 (4)" docSource="http://dx.doi.org/10.11646/phytotaxa.652.4.4" docStyle="DocumentStyle:96748F8F1B6C902996E134952A3A36B9.13:Phytotaxa.2014-.journal_article" docStyleId="96748F8F1B6C902996E134952A3A36B9" docStyleName="Phytotaxa.2014-.journal_article" docStyleVersion="13" docTitle="Tetralunata schoemanii, sp. nov." docType="treatment" docVersion="3" lastPageNumber="290" masterDocId="166DFFFCFFA4B075FF841F2EFF8E1E06" masterDocTitle="Light and Scanning Observations on a Denticula species reported from South Africa and endemism of the diatom genus Tetralunata (Rhopalodiales, Bacillariophyceae)" masterLastPageNumber="292" masterPageNumber="284" pageNumber="289" updateTime="1724864777837" updateUser="carolina" zenodo-license-figures="UNSPECIFIED" zenodo-license-treatments="UNSPECIFIED">
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<mods:title id="5E76F7FC8BAF606AB23E875B71B070A5">Light and Scanning Observations on a Denticula species reported from South Africa and endemism of the diatom genus Tetralunata (Rhopalodiales, Bacillariophyceae)</mods:title>
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<mods:namePart id="4D3641101E74572947E513AFEFAB7F0D">Kociolek, J. Patrick</mods:namePart>
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<mods:namePart id="877224C8AA13F716CD639EF471510952">Greenwood, Megan</mods:namePart>
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<mods:namePart id="73C869691264FB6E542B08CC7F7209D1">Rogers, Dillan</mods:namePart>
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<mods:namePart id="D2E98452D85B2C3323C3AAA5524EB572">Hamsher, Sarah E.</mods:namePart>
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<mods:affiliation id="13F270A89C338CF409BB97CC51D8705E">Department of Biology and Annis Water Resources Institute, Grand Valley State University, Allendale and Muskegon, Michigan, USA</mods:affiliation>
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<mods:affiliation id="C64B938237CEB4F515C0F3538291328D">Division of Biological Sciences, University of Montana, Missoula, Montana, USA 59812</mods:affiliation>
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<mods:namePart id="EBFE68587655AD389D24C09CD93F61F3">Li, Jingchun</mods:namePart>
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<mods:affiliation id="DEA9BBFA8B9E26B2425A7C4AE8A8691F">Museum of Natural History and Department of Ecology and Evolutionary Biology, University of Colorado, Boulder, CO 80309, USA</mods:affiliation>
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<mods:namePart id="4F7B7818D1CF840BD313CD0CC1B6E554">Chang, Aimee Caye G.</mods:namePart>
<mods:affiliation id="C18387FF003CC3B0541423DFA9CD914B">Museum of Natural History and Department of Ecology and Evolutionary Biology, University of Colorado, Boulder, CO 80309, USA &amp; Department of Biological Sciences, University of Santo Tomas, Sampaloc, Manila, Philippines</mods:affiliation>
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<mods:namePart id="FC1B2084D84F207BED0C4E9EE07BE6C6">Taylor, Jonathan</mods:namePart>
<mods:affiliation id="1CF20DD24270F5672E0434A6A366EFF3">South African National Diatom Collection and South African Institute for Aquatic Biodiversity (SAIAB), Makhanda, South Africa &amp; Unit for Environmental Science and Management, North-West University, Potchefstroom, South Africa</mods:affiliation>
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<mods:date id="C6FAA1ECBA23090E91641FD8B1989B9D">2024</mods:date>
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<taxonomicName id="A5FD4D11FFA1B070FF0C1AFBFE171BF6" authority="Kociolek et al." box="[136,409,1493,1520]" class="Bacillariophyceae" family="Rhopalodiaceae" genus="Tetralunata" higherTaxonomySource="GBIF" kingdom="Chromista" order="Rhopalodiales" pageId="5" pageNumber="289" phylum="Bacillariophyta" rank="species" species="schoemanii" status="sp. nov.">
<emphasis id="5089EA80FFA1B070FF0C1AFBFE171BF6" bold="true" box="[136,409,1493,1520]" italics="true" pageId="5" pageNumber="289">Tetralunata schoemanii</emphasis>
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<figureCitation id="FAC62A17FFA1B070FF0C1AD7FE951812" box="[136,283,1529,1556]" captionStart-0="FIGURES 110" captionStart-1="FIGURES 1115" captionStart-2="FIGURES 1619" captionStart-3="FIGURE 20, 21" captionStartId-0="2.[136,248,746,768]" captionStartId-1="3.[136,248,1953,1975]" captionStartId-2="4.[136,248,1953,1975]" captionStartId-3="5.[136,234,1101,1123]" captionTargetBox-0="[151,1435,190,722]" captionTargetBox-1="[213,1374,190,1928]" captionTargetBox-2="[278,1308,190,1929]" captionTargetBox-3="[151,1435,190,1077]" captionTargetId-0="figure-21@2.[151,1435,190,722]" captionTargetId-1="figure-34@3.[213,1374,190,1928]" captionTargetId-2="figure-21@4.[278,1308,190,1929]" captionTargetId-3="figure-34@5.[151,1435,190,1077]" captionTargetPageId-0="2" captionTargetPageId-1="3" captionTargetPageId-2="4" captionTargetPageId-3="5" captionText-0="FIGURES 110. Tetralunata schoemanii, sp. nov. Light Microscopy. Figs 18. Size diminution series, valve views, from Holotype slide. Fig. 9. Septa. Fig. 10. Frustule in girdle view showing Häutung valve. Scale bar = 10 µm for all specimens." captionText-1="FIGURES 1115. Tetralunata schoemanii, sp. nov. Scanning electron microscopy. Valve exterior. Figures 11, 12. Valve view of entire cell. Scale bars = 5 µm. Figs 13, 14. Higher magnification view of volae occluding areolae, on valve face and mantle, respectively. Scale bars = 1.5 µm. Fig. 15. Girdle view showing the girdle bands near the valve apex. Scale bar = 2.5 µm." captionText-2="FIGURES 1619. Tetralunata schoemanii, sp. nov. Scanning electron microscopy. Internal views. Fig. 16. Valve view with costae evident. Scale bar = 10 µm. Fig. 17. Valve with septum overlaying valve. Scale bar = 5 µm. Fig. 18. Septum. Scale bar = 5 µm. Fig. 19. High magnification view of the center of the valve showing proximal raphe ends. Scale bar = 1 µm." captionText-3="FIGURE 20, 21. Tetralunata schoemanii, sp. nov. Septum, cross section. Fig. 20. Septum only. Fig. 21. Valve with costa and septum overlaying the valve. Scale bars = 5 µm." figureDoi-0="http://doi.org/10.5281/zenodo.13216839" figureDoi-1="http://doi.org/10.5281/zenodo.13216841" figureDoi-2="http://doi.org/10.5281/zenodo.13216843" figureDoi-3="http://doi.org/10.5281/zenodo.13216845" httpUri-0="https://zenodo.org/record/13216839/files/figure.png" httpUri-1="https://zenodo.org/record/13216841/files/figure.png" httpUri-2="https://zenodo.org/record/13216843/files/figure.png" httpUri-3="https://zenodo.org/record/13216845/files/figure.png" pageId="5" pageNumber="289">Figures 121</figureCitation>
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<emphasis id="5089EA80FFA1B070FF0C196FFE92185D" bold="true" box="[136,284,1601,1627]" pageId="5" pageNumber="289">Description:</emphasis>
Valves are linear in shape with apices tapered and narrowly rounded. Length ranges from 2355 µm and breadth 710 µm. The raphe is placed medianly, with the raphe branches indistinct. The proximal ends terminate close to one another. Distally, the raphe branches extend onto the mantle at the apices. Costae are robust, 13 / 10 µm. Striae are parallel, composed of 46 areolae, and number 810 / 10 µm. Areolae clover-shaped with 48 c-shaped occlusions with those closer to the raphe having more occlusions and complexity. In girdle view, “capitate costae” are visible. Also evident in girdle view are Häutung valves, with a single new valve produced within a frustule. Häutung valves appear arched or undulate but otherwise are structured similar to vegetative valves.
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<emphasis id="5089EA80FFA1B070FF391810FE90195E" bold="true" box="[189,286,1854,1880]" pageId="5" pageNumber="289">Type:—</emphasis>
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treks camp, situated on the banks of the
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, on the road to Marakabei. Sides of hollow, cave-like cliffs below the road, damp rock faces. (
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! 652039, here illustrated as
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;
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! 82423,
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.
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<emphasis id="5089EA80FFA1B070FF3918E0FEEA19EE" bold="true" box="[189,356,1998,2024]" pageId="5" pageNumber="289">Etymology:—</emphasis>
Named in honor of F.R. Schoeman who collected and first reported this diatom.
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The production of Häutung valves as initially reported was thought to be unique to the genus
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</taxonomicName>
(
<bibRefCitation id="066C4B63FFA1B073FAE618DFFF4D1EBE" author="Thaler, F." lastPageId="6" lastPageNumber="290" pageId="5" pageNumber="289" pagination="313 - 347" refId="ref4919" refString="Thaler, F. (1972) Beitrag zur Entwicklungsgeschichte und zum Zellbau einiger Diatomeen. Osterreiche Botanische Zeitschrift 120: 313 - 347. https: // doi. org / 10.1007 / BF 01373382" type="journal article" year="1972">Thaler 1972</bibRefCitation>
;
<bibRefCitation id="066C4B63FFA2B073FF491FB0FECB1EBE" author="Sims, P. A." box="[205,325,158,184]" pageId="6" pageNumber="290" pagination="211 - 236" refId="ref4880" refString="Sims, P. A. (1983) A taxonomic study of the genus Epithemia with special reference to the type species E. turgida (Ehrenb.) Kutz. Bacillaria 6: 211 - 236." type="journal article" year="1983">Sims 1983</bibRefCitation>
), although Fricke (
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Schmidt) had illustrated it in
<taxonomicName id="A5FD4D11FFA2B073FC2E1FB0FB2F1EBE" authorityName="Brun" authorityYear="1891" box="[938,1185,158,184]" class="Bacillariophyceae" family="Bacillariaceae" genus="Denticula" higherTaxonomySource="GBIF" kingdom="Chromista" order="Bacillariales" pageId="6" pageNumber="290" phylum="Bacillariophyta" rank="species" species="vanheurckii">
<emphasis id="5089EA80FFA2B073FC2E1FB0FB2F1EBE" box="[938,1185,158,184]" italics="true" pageId="6" pageNumber="290">Denticula vanheurckii</emphasis>
</taxonomicName>
(=
<taxonomicName id="A5FD4D11FFA2B073FB3B1FB1FE261EDA" authority="(Brun) Hamsher et al. 2014: 361" authorityName="Hamsher" authorityPageNumber="361" authorityYear="2014" baseAuthorityName="Brun" class="Bacillariophyceae" family="Bacillariaceae" genus="Denticula" higherTaxonomySource="GBIF" kingdom="Chromista" order="Bacillariales" pageId="6" pageNumber="290" phylum="Bacillariophyta" rank="species" species="vanheurckii">
<emphasis id="5089EA80FFA2B073FB3B1FB1FAD51EBE" box="[1215,1371,158,184]" italics="true" pageId="6" pageNumber="290">T. vanheurckii</emphasis>
(Brun)
<bibRefCitation id="066C4B63FFA2B073FF0C1FECFE261EDA" author="Hamsher, S. E. &amp; Graeff, C. L. &amp; Stepanek, J. G. &amp; Kociolek, J. P." box="[136,424,194,220]" pageId="6" pageNumber="290" pagination="346 - 365" refId="ref3793" refString="Hamsher, S. E., Graeff, C. L., Stepanek, J. G. &amp; Kociolek, J. P. (2014) Variation in valve and girdle band morphology in freshwater Denticula (Bacillariophyceae) species: Implications for the systematic position of the genus including the description of Tetralunata gen. nov. (Epithemiaceae, Rhopalodiales). Plant Ecology and Evolution 147: 346 - 365. https: // doi. org / 10.5091 / plecevo. 2014.990" type="journal article" year="2014">
Hamsher
<emphasis id="5089EA80FFA2B073FF701FEDFEA31EDA" box="[244,301,194,220]" italics="true" pageId="6" pageNumber="290">et al.</emphasis>
2014: 361
</bibRefCitation>
</taxonomicName>
). Häutung valves were first described by
<bibRefCitation id="066C4B63FFA2B073FCFD1FECFB971EDA" author="Geitler, L." box="[889,1049,194,220]" pageId="6" pageNumber="290" pagination="98 - 100" refId="ref3594" refString="Geitler, L. (1927) Hautung bei einer pennaten Diatomee. Osterreiche Botanische Zeitschrift 76: 98 - 100. https: // doi. org / 10.1007 / BF 01246242" type="journal article" year="1927">Geitler (1927)</bibRefCitation>
and later in greater detail by
<bibRefCitation id="066C4B63FFA2B073FAE61FECFF5D1F06" author="Thaler, F." pageId="6" pageNumber="290" pagination="313 - 347" refId="ref4919" refString="Thaler, F. (1972) Beitrag zur Entwicklungsgeschichte und zum Zellbau einiger Diatomeen. Osterreiche Botanische Zeitschrift 120: 313 - 347. https: // doi. org / 10.1007 / BF 01373382" type="journal article" year="1972">Thaler (1972)</bibRefCitation>
as a single valve produced within a frustule that more or less resembles the vegetative valves, distinguishing it from Innenschalen (e.g.
<bibRefCitation id="066C4B63FFA2B073FE351E24FDCF1F22" author="Geitler, L." box="[433,577,266,292]" pageId="6" pageNumber="290" pagination="275 - 286" refId="ref3672" refString="Geitler, L. (1980) Zellteilung und Bildung von Innenschalen bei Hantzschia amphioxys und Achnanthes coarctata. Plant Systematics and Evolution 136: 275 - 286. https: // doi. org / 10.1007 / BF 01004632" type="journal article" year="1980">Geitler 1980</bibRefCitation>
;
<bibRefCitation id="066C4B63FFA2B073FDC91E24FCBD1F22" author="Kociolek, J. P. &amp; Liu, Y. &amp; Wang, Q. X." box="[589,819,266,292]" pageId="6" pageNumber="290" pagination="486 - 494" refId="ref4209" refString="Kociolek, J. P., Liu, Y. &amp; Wang, Q. X. (2011) Internal valves in populations of Meridion circulare from the A'er Mountain region of northeasern China: Implications for taxonomy and systematics. Journal of Systematics and Evolution 49 (5): 486 - 494. https: // doi. org / 10.1111 / j. 1759 - 6831.2011.00142. x" type="journal article" year="2011">
Kociolek
<emphasis id="5089EA80FFA2B073FD3E1E25FD7D1F22" box="[698,755,266,292]" italics="true" pageId="6" pageNumber="290">et al.</emphasis>
2011
</bibRefCitation>
) and valves produced within cells of
<taxonomicName id="A5FD4D11FFA2B073FB531E25FF731F4E" authority="Ehrenberg (1837: 44)" authorityName="Ehrenberg" authorityPageNumber="44" authorityYear="1837" class="Bacillariophyceae" family="Eunotiaceae" genus="Eunotia" higherTaxonomySource="GBIF" kingdom="Chromista" order="Eunotiales" pageId="6" pageNumber="290" phylum="Bacillariophyta" rank="genus">
<emphasis id="5089EA80FFA2B073FB531E25FABE1F22" box="[1239,1328,267,292]" italics="true" pageId="6" pageNumber="290">Eunotia</emphasis>
<bibRefCitation id="066C4B63FFA2B073FAB31E24FF731F4E" author="Ehrenberg, C. G." pageId="6" pageNumber="290" pagination="43 - 45" refId="ref3484" refString="Ehrenberg, C. G. (1837) Uber ein aus fossilen Infusorien bestehendes, 1832 zu Brod verbacknes Bergmehl von der Grenzen Lapplands in Schweden. Bericht uber die zur Bekanntmachung geeigneten Verhandlungen der Koniglich-Preussischen Akademie der Wissenschaften zu Berlin 1837: 43 - 45. https: // doi. org / 10.1002 / andp. 18371160116" type="journal article" year="1837">Ehrenberg (1837: 44)</bibRefCitation>
</taxonomicName>
(e.g.
<bibRefCitation id="066C4B63FFA2B073FEB81E01FDE21F4E" author="Von Stosch, H. - A. &amp; Fecher, K." box="[316,620,302,329]" pageId="6" pageNumber="290" pagination="233 - 243" refId="ref4955" refString="Von Stosch, H. - A. &amp; Fecher, K. (1979) &quot; Internal thecae &quot; of Eunotia soleirolii (Bacillariophyceae): development, structure and function as resting spores. Journal of Phycology 15 (3): 233 - 243. https: // doi. org / 10.1111 / j. 0022 - 3646.1979.00233. x" type="journal article" year="1979">Von Stosch &amp; Fecher 1979</bibRefCitation>
). These valves may have undulate valve faces, making them distinct in the light microscope. The reason for their production is uncertain, but they may be reactions to osmostic stress or serve as a resting stage (
<bibRefCitation id="066C4B63FFA2B073FEBD1E58FE4B1F96" author="Thaler, F." box="[313,453,374,400]" pageId="6" pageNumber="290" pagination="313 - 347" refId="ref4919" refString="Thaler, F. (1972) Beitrag zur Entwicklungsgeschichte und zum Zellbau einiger Diatomeen. Osterreiche Botanische Zeitschrift 120: 313 - 347. https: // doi. org / 10.1007 / BF 01373382" type="journal article" year="1972">Thaler 1972</bibRefCitation>
).
</paragraph>
<paragraph id="62423692FFA2B073FF391EB4FC191C42" blockId="6.[136,1452,158,833]" pageId="6" pageNumber="290">
Observations on the
<taxonomicName id="A5FD4D11FFA2B073FE2A1EB4FD051FB2" box="[430,651,410,436]" pageId="6" pageNumber="290">
<emphasis id="5089EA80FFA2B073FE2A1EB4FDBE1FB2" box="[430,560,410,436]" italics="true" pageId="6" pageNumber="290">Tetralunata</emphasis>
species
</taxonomicName>
from
<collectingCountry id="1AEA7602FFA2B073FD521EB4FCE61FB2" box="[726,872,410,436]" name="South Africa" pageId="6" pageNumber="290">South Africa</collectingCountry>
confirms their close relationship to
<taxonomicName id="A5FD4D11FFA2B073FA941EB4FA0D1FB2" box="[1296,1411,410,436]" class="Bacillariophyceae" family="Rhopalodiaceae" genus="Epithemia" higherTaxonomySource="GBIF" kingdom="Chromista" order="Rhopalodiales" pageId="6" pageNumber="290" phylum="Bacillariophyta" rank="genus">
<emphasis id="5089EA80FFA2B073FA941EB4FA0D1FB2" box="[1296,1411,410,436]" italics="true" pageId="6" pageNumber="290">Epithemia</emphasis>
</taxonomicName>
, as suggested by
<bibRefCitation id="066C4B63FFA2B073FEA71E90FDAF1FDE" author="Hamsher, S. E. &amp; Graeff, C. L. &amp; Stepanek, J. G. &amp; Kociolek, J. P." box="[291,545,446,472]" pageId="6" pageNumber="290" pagination="346 - 365" refId="ref3793" refString="Hamsher, S. E., Graeff, C. L., Stepanek, J. G. &amp; Kociolek, J. P. (2014) Variation in valve and girdle band morphology in freshwater Denticula (Bacillariophyceae) species: Implications for the systematic position of the genus including the description of Tetralunata gen. nov. (Epithemiaceae, Rhopalodiales). Plant Ecology and Evolution 147: 346 - 365. https: // doi. org / 10.5091 / plecevo. 2014.990" type="journal article" year="2014">
Hamsher
<emphasis id="5089EA80FFA2B073FE151E91FE451FDE" box="[401,459,446,472]" italics="true" pageId="6" pageNumber="290">et al.</emphasis>
(2014)
</bibRefCitation>
. Evidence for a close relationship to
<taxonomicName id="A5FD4D11FFA2B073FC4C1E90FBB51FDE" box="[968,1083,446,472]" class="Bacillariophyceae" family="Rhopalodiaceae" genus="Epithemia" higherTaxonomySource="GBIF" kingdom="Chromista" order="Rhopalodiales" pageId="6" pageNumber="290" phylum="Bacillariophyta" rank="genus">
<emphasis id="5089EA80FFA2B073FC4C1E90FBB51FDE" box="[968,1083,446,472]" italics="true" pageId="6" pageNumber="290">Epithemia</emphasis>
</taxonomicName>
is evidenced with respect to the presence and similarity in structure in the volae, septa, septa that clasp the costate fibulae forming capitate costae, lack of a keel, no poroids in the girdle bands, frustules that are planar and not segment-like in their organization, and production of Häutung valves (
<bibRefCitation id="066C4B63FFA2B073FE601D04FDEE1C42" author="Sims, P. A." box="[484,608,554,580]" pageId="6" pageNumber="290" pagination="211 - 236" refId="ref4880" refString="Sims, P. A. (1983) A taxonomic study of the genus Epithemia with special reference to the type species E. turgida (Ehrenb.) Kutz. Bacillaria 6: 211 - 236." type="journal article" year="1983">Sims 1983</bibRefCitation>
, Kociolek
<emphasis id="5089EA80FFA2B073FD5D1D05FC9F1C42" box="[729,785,554,580]" italics="true" pageId="6" pageNumber="290">et al.</emphasis>
submitted).
</paragraph>
<paragraph id="62423692FFA2B073FF391D61FDDD1D47" blockId="6.[136,1452,158,833]" pageId="6" pageNumber="290">
In terms of biogeography,
<bibRefCitation id="066C4B63FFA2B073FE681D60FD281C6E" author="Kociolek, J. P." box="[492,678,590,616]" pageId="6" pageNumber="290" pagination="509 - 534" refId="ref4165" refString="Kociolek, J. P. (2019) A worldwide listing and biogeography of freshwater diatom genera: A phylogenetic perspective. Diatom Research 33: 509 - 534. https: // doi. org / 10.1080 / 0269249 X. 2019.1574243" type="journal article" year="2019">Kociolek (2019)</bibRefCitation>
had indicated
<taxonomicName id="A5FD4D11FFA2B073FCCB1D60FC5F1C6E" box="[847,977,590,616]" class="Bacillariophyceae" family="Rhopalodiaceae" genus="Tetralunata" higherTaxonomySource="GBIF" kingdom="Chromista" order="Rhopalodiales" pageId="6" pageNumber="290" phylum="Bacillariophyta" rank="genus">
<emphasis id="5089EA80FFA2B073FCCB1D60FC5F1C6E" box="[847,977,590,616]" italics="true" pageId="6" pageNumber="290">Tetralunata</emphasis>
</taxonomicName>
was endemic to
<collectingCountry id="1AEA7602FFA2B073FB101D60FA8B1C6E" box="[1172,1285,590,616]" name="Indonesia" pageId="6" pageNumber="290">Indonesia</collectingCountry>
. Reporting the genus in
<collectingCountry id="1AEA7602FFA2B073FF6A1D5CFEF31C8A" box="[238,381,626,652]" name="South Africa" pageId="6" pageNumber="290">South Africa</collectingCountry>
here for the first time, as a species endemic to southern Africa, expands the known distribution of the genus considerably. Our current understanding is that the members of the
<taxonomicName id="A5FD4D11FFA2B073FBA31DB8FB491CB6" box="[1063,1223,662,688]" class="Bacillariophyceae" higherTaxonomySource="GBIF" kingdom="Chromista" order="Rhopalodiales" pageId="6" pageNumber="290" phylum="Bacillariophyta" rank="order">Rhopalodiales</taxonomicName>
have verified fossil records that extend back 30 mya (
<bibRefCitation id="066C4B63FFA2B073FD8D1D94FD6B1CD2" author="Benson, M. E. &amp; Kociolek, J. P. &amp; Spaulding, S. A. &amp; Smith, D. M." box="[521,741,698,724]" pageId="6" pageNumber="290" pagination="131 - 152" refId="ref3359" refString="Benson, M. E., Kociolek, J. P., Spaulding, S. A. &amp; Smith, D. M. (2012) Pre-Neogene (&gt; 23.8 MA) biochronology of non-marine diatoms, with an account of the late Eocene flora of the Florissant Formation, central Colorado, USA. Stratigraphy 9: 131 - 152. https: // doi. org / 10.29041 / strat. 09.2.02" type="journal article" year="2012">
Benson
<emphasis id="5089EA80FFA2B073FDE21D95FD2E1CD2" box="[614,672,698,724]" italics="true" pageId="6" pageNumber="290">et al.</emphasis>
2012
</bibRefCitation>
). Given the wide separation in both space and time of SE Asia and
<collectingCountry id="1AEA7602FFA2B073FF331DF0FECB1CFE" box="[183,325,734,760]" name="South Africa" pageId="6" pageNumber="290">South Africa</collectingCountry>
during the breakup of the continents (
<bibRefCitation id="066C4B63FFA2B073FD6E1DF0FC341CFE" author="McLoughlin, S." box="[746,954,734,760]" pageId="6" pageNumber="290" pagination="271 - 300" refId="ref4596" refString="McLoughlin, S. (2001) The breakup history of Gondwana and its impact on pre-Cenozoic floristic provincialism. Australian Journal of Botany 49 (3): 271 - 300. https: // doi. org / 10.1071 / BT 00023" type="journal article" year="2001">McLoughlin 2001</bibRefCitation>
), earth history dynamics would not be useful to identify a mechanism for this disjunct distribution. Further research will be necessary to attain an understanding of the separate locations for this rare genus.
</paragraph>
</subSubSection>
</treatment>
</document>