291 lines
57 KiB
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291 lines
57 KiB
XML
<document id="D3D5B165AF52F4C8470C10C27A7415A8" ID-DOI="10.4467/16890027AP.14.022.1997" ID-ISSN="1689-0027" ID-Zenodo-Dep="12638350" IM.bibliography_approvedBy="felipe" IM.illustrations_approvedBy="felipe" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="GgImagineBatch" IM.taxonomicNames_approvedBy="felipe" IM.treatments_approvedBy="felipe" checkinTime="1720037563826" checkinUser="felipe" docAuthor="Thomsen, Helge A. & Østergaard, Jette B." docDate="2014" docId="AC4A6B03FF8EFFF4FCECD869FADDFA9E" docLanguage="en" docName="ActaProtozool.53.3.235-256.pdf" docOrigin="Acta Protozoologica 53 (3)" docSource="https://www.mendeley.com/catalogue/4d144474-225a-3934-8814-0d8c1ebb76df/" docStyle="DocumentStyle:64ABA4BEA387EE5C24A0DDEB32CC92E7.3:ActaProtozool.2014-.journal_article" docStyleId="64ABA4BEA387EE5C24A0DDEB32CC92E7" docStyleName="ActaProtozool.2014-.journal_article" docStyleVersion="3" docTitle="Pappomonas flabellifera Manton and Oates 1975" docType="treatment" docVersion="2" lastPageNumber="239" masterDocId="5073137BFF8CFFF0FFDCD910FFDDFFAD" masterDocTitle="Coccolithophorids in Polar Waters: Pappomonas spp. Revisited" masterLastPageNumber="256" masterPageNumber="235" pageNumber="237" updateTime="1720040505996" updateUser="ExternalLinkService" zenodo-license-document="CC-BY-4.0">
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<mods:title id="BA26CDA9C417D5D74B5FB3220BFD5DDC">Coccolithophorids in Polar Waters: Pappomonas spp. Revisited</mods:title>
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<paragraph id="245CDA15FF8EFFF2FCECD869FA56FE3E" blockId="2.[816,1419,377,404]" box="[816,1419,377,404]" pageId="2" pageNumber="237">
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<heading id="7F146D79FF8EFFF2FCECD869FA56FE3E" box="[816,1419,377,404]" fontSize="11" level="2" pageId="2" pageNumber="237" reason="5">
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<taxonomicName id="E3E3A196FF8EFFF2FCECD869FA56FE3E" authority="Manton and Oates 1975" authorityName="Manton and Oates" authorityYear="1975" box="[816,1419,377,404]" class="Coccolithophyceae" family="Papposphaeraceae" genus="Pappomonas" kingdom="Chromista" order="Coccosphaerales" pageId="2" pageNumber="237" phylum="Haptophyta" rank="species" species="flabellifera">
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<emphasis id="16970607FF8EFFF2FCECD869FA56FE3E" bold="true" box="[816,1419,377,404]" pageId="2" pageNumber="237">
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<emphasis id="16970607FF8EFFF2FCECD869FB85FE39" bold="true" box="[816,1112,377,404]" italics="true" pageId="2" pageNumber="237">Pappomonas flabellifera</emphasis>
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<bibRefCitation id="4072A7E4FF8EFFF2FB83D869FA56FE3E" author="Manton I. & Oates K." box="[1119,1419,377,403]" pageId="2" pageNumber="237" pagination="93 - 109" refId="ref9975" refString="Manton I., Oates K. (1975) Fine-structural observations on Papposphaera Tangen from the southern hemisphere and on Pappomonas gen. nov. from South Africa and Greenland. Brit. Phycol. J. 10: 93 - 109" type="journal article" year="1975">Manton and Oates 1975</bibRefCitation>
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</emphasis>
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</taxonomicName>
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</heading>
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</paragraph>
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</subSubSection>
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<subSubSection id="6CF9899EFF8EFFF4FC8AD8BBFADDFA9E" lastPageId="4" lastPageNumber="239" pageId="2" pageNumber="237" type="materials_examined">
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<paragraph id="245CDA15FF8EFFF2FC8AD8BBFAC2FAE7" blockId="2.[816,1463,427,1944]" pageId="2" pageNumber="237">
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When first described (
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<bibRefCitation id="4072A7E4FF8EFFF2FB82D8BBFA5CFE6B" author="Manton I. & Oates K." box="[1118,1409,427,454]" pageId="2" pageNumber="237" pagination="93 - 109" refId="ref9975" refString="Manton I., Oates K. (1975) Fine-structural observations on Papposphaera Tangen from the southern hemisphere and on Pappomonas gen. nov. from South Africa and Greenland. Brit. Phycol. J. 10: 93 - 109" type="journal article" year="1975">Manton and Oates 1975</bibRefCitation>
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) the material examined comprised as also acknowledged by the authors three morphotypes that were basically dis- tinguished by differences in the appearance of the blades that terminate the circumflagellar coccoliths. While the West
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<collectingCountry id="5CF49A85FF8EFFF2FCADDB48FC36FDDF" box="[881,1003,600,626]" name="Greenland" pageId="2" pageNumber="237">Greenland</collectingCountry>
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morphotype (loc.cit.
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<figureCitation id="BCD8C690FF8EFFF2FB3ADB48FA5AFDDF" box="[1254,1415,600,626]" captionStart="Figs 9–17" captionStartId="8.[132,175,1752,1774]" captionText="Figs 9–17. Pappomonas flabellifera from Svalbard (9–12), Mexico (13–14) and California (15–17). SEM (9–12) and TEM (13–17) whole mounts. 9 – complete cell showing closely packed body coccoliths and a very well defined ring of circumflagellar coccoliths (ca. 15); 10 – detail of body coccoliths showing the upright position of rim elements; 11 – complete cell; 12 – detail of circumflagellar coccoliths (from Fig. 9) indicating that neighbouring ‘half blades’ of a triangle are joined at a blunt angle; 13 – whole cell very reminiscent of material previously published from S. Africa; see text for further details; 14 – detail of a reduced central appendage formed by two equally sized half blades; 15 – whole cell with flagella and haptonema; 16 – detail from Fig. 15 showing triangles that are reminiscent of those of the S.African type cell; see text for further details; 17 – detail from Fig. 15 of reduced, yet stalked central appendages." pageId="2" pageNumber="237">Figs 9–12, 14</figureCitation>
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,
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<figureCitation id="BCD8C690FF8EFFF2FA4EDB48FC93FD38" captionStart="Figs 19–26" captionStartId="10.[134,177,1716,1738]" captionText="Figs 19–26. Pappomonas borealis from the Arctic (NEW / Figs 19–24; Svalbard / Figs 25–26); TEM (Figs 19–24) and SEM whole mounts (Figs 25–26). 19 – single body coccolith (reversed printing / detail from Fig. 20) showing rim (pentagonals and rods) as well as central area calcification consisting of two ovals and four embedded lines of rod-shaped elements in the middle; notice that the radiating fibrils on the upper surface of the base plate scale are visible in between elements of the central area calcification; 20 – whole cell with flagella and a curled up haptonema; 21 – detail of posterior cell end body coccoliths from the cell shown in Fig. 20; 22 – detail of body coccoliths from the cell shown in Fig. 23; 23 – whole cell; 24 – detail of circumflagellar coccoliths; notice the extended overlap between shaft and triangu- lar blade; 25 – detail of anterior cell end to show the flattened triangular blade and the upright rim on body coccoliths; 26 – a sequence of circumflagellar coccoliths showing the upright rim and the cruciform central area calcification." pageId="2" pageNumber="237">20, 21</figureCitation>
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) was later singled out as the variety
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<taxonomicName id="E3E3A196FF8EFFF2FB2ADB6BFA82FD38" authority="(Manton et al. 1976a)" authorityName="Thomsen & Østergaard" authorityYear="2014" baseAuthorityName="Manton, Sutherland and McCully" baseAuthorityYear="1976" box="[1270,1375,635,661]" class="Coccolithophyceae" family="Papposphaeraceae" genus="Pappomonas" kingdom="Chromista" order="Coccosphaerales" pageId="2" pageNumber="237" phylum="Haptophyta" rank="species" species="borealis">
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‘
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<emphasis id="16970607FF8EFFF2FADCDB6BFA82FD38" box="[1280,1375,635,661]" italics="true" pageId="2" pageNumber="237">borealis</emphasis>
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</taxonomicName>
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’ (
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<bibRefCitation id="4072A7E4FF8EFFF2FAA9DB6BFC30FD1A" author="Manton I. & Sutherland J. & McCully M." pageId="2" pageNumber="237" pagination="225 - 238" refId="ref10063" refString="Manton I., Sutherland J., McCully M. (1976 a) Fine structural observations on coccolithophorids from South Alaska in the genera Papposphaera Tangen and Pappomonas Manton et Oates. Brit. Phycol. J. 11: 225 - 238" type="journal article" year="1976">
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Manton
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<emphasis id="16970607FF8EFFF2FC82DB8EFC44FD15" box="[862,921,670,696]" italics="true" pageId="2" pageNumber="237">et al.</emphasis>
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1976a
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</bibRefCitation>
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), the two remaining morphotypes both of S. African origin were maintained within
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<taxonomicName id="E3E3A196FF8EFFF2FAE2DBD1FBCAFD50" authority="var. flabellifera" class="Coccolithophyceae" family="Papposphaeraceae" genus="Pappomonas" kingdom="Chromista" order="Coccosphaerales" pageId="2" pageNumber="237" phylum="Haptophyta" rank="variety" species="flabellifera" variety="flabellifera">
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<emphasis id="16970607FF8EFFF2FAE2DBD1FC82FD50" italics="true" pageId="2" pageNumber="237">P. flabellifera</emphasis>
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var.
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<emphasis id="16970607FF8EFFF2FC4ADBF3FBCAFD50" box="[918,1047,739,765]" italics="true" pageId="2" pageNumber="237">flabellifera</emphasis>
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</taxonomicName>
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. The S.African
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<typeStatus id="FB5864B7FF8EFFF2FB0CDBF3FADEFD50" box="[1232,1283,739,765]" pageId="2" pageNumber="237">type</typeStatus>
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material (
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<bibRefCitation id="4072A7E4FF8EFFF2FAA9DBF3FBC6FC8D" author="Manton I. & Oates K." pageId="2" pageNumber="237" pagination="93 - 109" refId="ref9975" refString="Manton I., Oates K. (1975) Fine-structural observations on Papposphaera Tangen from the southern hemisphere and on Pappomonas gen. nov. from South Africa and Greenland. Brit. Phycol. J. 10: 93 - 109" type="journal article" year="1975">Manton and Oates 1975</bibRefCitation>
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; loc.cit.
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<figureCitation id="BCD8C690FF8EFFF2FB5CDA16FB34FCB2" box="[1152,1257,773,800]" captionStart="Figs 3–8" captionStartId="4.[134,177,1749,1771]" captionText="Figs 3–8. Pappomonas flabellifera TEM whole mounts from the Arctic (NEW). 3 – whole cell with details accounted for in Figs 6–7; the arrow points to the demarcation line between the significantly different sized half blades of the triangle; 4 – whole cell with flagella and a curled up haptonema; 5 – detail from Fig. 4 showing body coccoliths with a fairly extensive central area calcification; reversed printing; 6 – body coccoliths (detail from Fig. 3) showing rim and central area calcification; 7 – oval flagellar pole coccolith with extensive central area calcification; notice the minute elements making up the stalk and the short overlap between stalk and triangle (arrows); 8 – detached armour of coccoliths; circumflagellar coccoliths (arrows) are oval, nearly of the same size as body coccoliths and with fairly extensive calcification." pageId="2" pageNumber="237">Figs 5, 6</figureCitation>
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) which is further substantiated by additional micrographs (loc.cit.
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<figureCitation id="BCD8C690FF8EFFF2FA58DA38FC82FCC8" captionStart="Figs 3–8" captionStartId="4.[134,177,1749,1771]" captionText="Figs 3–8. Pappomonas flabellifera TEM whole mounts from the Arctic (NEW). 3 – whole cell with details accounted for in Figs 6–7; the arrow points to the demarcation line between the significantly different sized half blades of the triangle; 4 – whole cell with flagella and a curled up haptonema; 5 – detail from Fig. 4 showing body coccoliths with a fairly extensive central area calcification; reversed printing; 6 – body coccoliths (detail from Fig. 3) showing rim and central area calcification; 7 – oval flagellar pole coccolith with extensive central area calcification; notice the minute elements making up the stalk and the short overlap between stalk and triangle (arrows); 8 – detached armour of coccoliths; circumflagellar coccoliths (arrows) are oval, nearly of the same size as body coccoliths and with fairly extensive calcification." pageId="2" pageNumber="237">Figs 7, 8</figureCitation>
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,
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<figureCitation id="BCD8C690FF8EFFF2FCB2DA5BFC64FCC8" box="[878,953,843,869]" captionStart="Figs 9–17" captionStartId="8.[132,175,1752,1774]" captionText="Figs 9–17. Pappomonas flabellifera from Svalbard (9–12), Mexico (13–14) and California (15–17). SEM (9–12) and TEM (13–17) whole mounts. 9 – complete cell showing closely packed body coccoliths and a very well defined ring of circumflagellar coccoliths (ca. 15); 10 – detail of body coccoliths showing the upright position of rim elements; 11 – complete cell; 12 – detail of circumflagellar coccoliths (from Fig. 9) indicating that neighbouring ‘half blades’ of a triangle are joined at a blunt angle; 13 – whole cell very reminiscent of material previously published from S. Africa; see text for further details; 14 – detail of a reduced central appendage formed by two equally sized half blades; 15 – whole cell with flagella and haptonema; 16 – detail from Fig. 15 showing triangles that are reminiscent of those of the S.African type cell; see text for further details; 17 – detail from Fig. 15 of reduced, yet stalked central appendages." pageId="2" pageNumber="237">16, 17</figureCitation>
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) represents a morphotype of
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<taxonomicName id="E3E3A196FF8EFFF2FAC4DA5CFA6AFCC8" authorityName="Manton and Oates" authorityYear="1975" box="[1304,1463,843,869]" class="Coccolithophyceae" family="Papposphaeraceae" genus="Pappomonas" kingdom="Chromista" order="Coccosphaerales" pageId="2" pageNumber="237" phylum="Haptophyta" rank="species" species="flabellifera">
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<emphasis id="16970607FF8EFFF2FAC4DA5CFA6AFCC8" box="[1304,1463,843,869]" italics="true" pageId="2" pageNumber="237">P. flabellifera</emphasis>
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</taxonomicName>
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that is rarely encountered and to the best of our knowledge not illustrated in any subsequent publication. The feature that most clearly identifies the
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<typeStatus id="FB5864B7FF8EFFF2FB22DAA3FAECFC60" box="[1278,1329,947,973]" pageId="2" pageNumber="237">type</typeStatus>
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material is the V-shaped circumflagellar coccolith appendage. The second S. African morphotype (loc.cit.
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<figureCitation id="BCD8C690FF8EFFF2FB23DAE8FA58FBBE" box="[1279,1413,1016,1043]" captionStart="Figs 9–17" captionStartId="8.[132,175,1752,1774]" captionText="Figs 9–17. Pappomonas flabellifera from Svalbard (9–12), Mexico (13–14) and California (15–17). SEM (9–12) and TEM (13–17) whole mounts. 9 – complete cell showing closely packed body coccoliths and a very well defined ring of circumflagellar coccoliths (ca. 15); 10 – detail of body coccoliths showing the upright position of rim elements; 11 – complete cell; 12 – detail of circumflagellar coccoliths (from Fig. 9) indicating that neighbouring ‘half blades’ of a triangle are joined at a blunt angle; 13 – whole cell very reminiscent of material previously published from S. Africa; see text for further details; 14 – detail of a reduced central appendage formed by two equally sized half blades; 15 – whole cell with flagella and haptonema; 16 – detail from Fig. 15 showing triangles that are reminiscent of those of the S.African type cell; see text for further details; 17 – detail from Fig. 15 of reduced, yet stalked central appendages." pageId="2" pageNumber="237">Figs 13, 15</figureCitation>
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,
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<figureCitation id="BCD8C690FF8EFFF2FA4EDAE8FA6CFBBF" box="[1426,1457,1016,1042]" captionStart="Fig" captionStartId="8.[816,851,743,765]" captionTargetBox="[829,1453,240,697]" captionTargetId="figure-476@8.[816,1466,223,712]" captionTargetPageId="8" captionText="Fig. 18. Pappomonas flabellifera (squares) and P. borealis (triangles) body scale dimensions." figureDoi="http://doi.org/10.5281/zenodo.12638354" httpUri="https://zenodo.org/record/12638354/files/figure.png" pageId="2" pageNumber="237">18</figureCitation>
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,
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<figureCitation id="BCD8C690FF8EFFF2FCECDD0BFC93FB98" box="[816,846,1051,1077]" captionStart="Figs 19–26" captionStartId="10.[134,177,1716,1738]" captionText="Figs 19–26. Pappomonas borealis from the Arctic (NEW / Figs 19–24; Svalbard / Figs 25–26); TEM (Figs 19–24) and SEM whole mounts (Figs 25–26). 19 – single body coccolith (reversed printing / detail from Fig. 20) showing rim (pentagonals and rods) as well as central area calcification consisting of two ovals and four embedded lines of rod-shaped elements in the middle; notice that the radiating fibrils on the upper surface of the base plate scale are visible in between elements of the central area calcification; 20 – whole cell with flagella and a curled up haptonema; 21 – detail of posterior cell end body coccoliths from the cell shown in Fig. 20; 22 – detail of body coccoliths from the cell shown in Fig. 23; 23 – whole cell; 24 – detail of circumflagellar coccoliths; notice the extended overlap between shaft and triangu- lar blade; 25 – detail of anterior cell end to show the flattened triangular blade and the upright rim on body coccoliths; 26 – a sequence of circumflagellar coccoliths showing the upright rim and the cruciform central area calcification." pageId="2" pageNumber="237">19</figureCitation>
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) has, however, become the de facto
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<taxonomicName id="E3E3A196FF8EFFF2FAC9DD0CFC3EFBF5" authority="var. flabellifera" class="Coccolithophyceae" family="Papposphaeraceae" genus="Pappomonas" kingdom="Chromista" order="Coccosphaerales" pageId="2" pageNumber="237" phylum="Haptophyta" rank="variety" species="flabellifera" variety="flabellifera">
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<emphasis id="16970607FF8EFFF2FAC9DD0CFA6AFB98" box="[1301,1463,1051,1077]" italics="true" pageId="2" pageNumber="237">P. flabellifera</emphasis>
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var.
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<emphasis id="16970607FF8EFFF2FCBEDD2EFC3EFBF5" box="[866,995,1086,1112]" italics="true" pageId="2" pageNumber="237">flabellifera</emphasis>
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</taxonomicName>
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which has later been found and reported from a range of geographic sites. The inclusion of no less than three morphotypes in the original description of
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<taxonomicName id="E3E3A196FF8EFFF2FC8FDDB6FC29FB6D" authorityName="Manton and Oates" authorityYear="1975" box="[851,1012,1190,1216]" class="Coccolithophyceae" family="Papposphaeraceae" genus="Pappomonas" kingdom="Chromista" order="Coccosphaerales" pageId="2" pageNumber="237" phylum="Haptophyta" rank="species" species="flabellifera">
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<emphasis id="16970607FF8EFFF2FC8FDDB6FC29FB6D" box="[851,1012,1190,1216]" italics="true" pageId="2" pageNumber="237">P. flabellifera</emphasis>
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</taxonomicName>
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has created confusion in as much as the diagnosis as it stands (and despite the precise identification of a
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<typeStatus id="FB5864B7FF8EFFF2FC0ADDFBFBD4FAA8" box="[982,1033,1259,1285]" pageId="2" pageNumber="237">type</typeStatus>
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specimen) does in fact include morphological features and dimensions sampled across the whole range of morphotypes encountered.
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</paragraph>
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<paragraph id="245CDA15FF8EFFF2FC8ADC43FC64F9B7" blockId="2.[816,1463,427,1944]" pageId="2" pageNumber="237">
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The below reexamination of a substantially larger material of
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<taxonomicName id="E3E3A196FF8EFFF2FC1ADC66FA6AFA3D" authority="var. flabellifera" box="[966,1463,1398,1424]" class="Coccolithophyceae" family="Papposphaeraceae" genus="Pappomonas" kingdom="Chromista" order="Coccosphaerales" pageId="2" pageNumber="237" phylum="Haptophyta" rank="variety" species="flabellifera" variety="flabellifera">
|
||
<emphasis id="16970607FF8EFFF2FC1ADC66FB33FA3D" box="[966,1262,1398,1424]" italics="true" pageId="2" pageNumber="237">Pappomonas flabellifera</emphasis>
|
||
var.
|
||
<emphasis id="16970607FF8EFFF2FAEADC66FA6AFA3D" box="[1334,1463,1398,1424]" italics="true" pageId="2" pageNumber="237">flabellifera</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="E3E3A196FF8EFFF2FCB6DC89FB67FA1F" box="[874,1210,1432,1459]" class="Coccolithophyceae" family="Papposphaeraceae" genus="Pappomonas" kingdom="Chromista" order="Coccosphaerales" pageId="2" pageNumber="237" phylum="Haptophyta" rank="variety" species="flabellifera" variety="borealis">
|
||
<emphasis id="16970607FF8EFFF2FCB6DC89FBCCFA1F" box="[874,1041,1432,1458]" italics="true" pageId="2" pageNumber="237">P. flabellifera</emphasis>
|
||
var.
|
||
<emphasis id="16970607FF8EFFF2FB87DC88FB67FA1F" box="[1115,1210,1432,1458]" italics="true" pageId="2" pageNumber="237">borealis</emphasis>
|
||
</taxonomicName>
|
||
(under the heading
|
||
<taxonomicName id="E3E3A196FF8EFFF2FCECDCACFC6DFA78" authorityName="Thomsen & Østergaard" authorityYear="2014" baseAuthorityName="Manton, Sutherland and McCully" baseAuthorityYear="1976" box="[816,944,1467,1493]" class="Coccolithophyceae" family="Papposphaeraceae" genus="Pappomonas" kingdom="Chromista" order="Coccosphaerales" pageId="2" pageNumber="237" phylum="Haptophyta" rank="species" species="borealis" status="stat. nov.">
|
||
<emphasis id="16970607FF8EFFF2FCECDCACFC6DFA78" box="[816,944,1467,1493]" italics="true" pageId="2" pageNumber="237">P. borealis</emphasis>
|
||
</taxonomicName>
|
||
<taxonomicNameLabel id="0DA4BB7CFF8EFFF2FC67DCABFBF7FA78" box="[955,1066,1467,1493]" pageId="2" pageNumber="237">stat. nov.</taxonomicNameLabel>
|
||
) provides in our opinion the in- sight needed to set up a more robust future taxonomic framework.
|
||
</paragraph>
|
||
<paragraph id="245CDA15FF8EFFF2FC8ADF33FA7EF908" blockId="2.[816,1463,427,1944]" pageId="2" pageNumber="237">
|
||
The vast majority of the
|
||
<taxonomicName id="E3E3A196FF8EFFF2FB58DF34FAF9F990" authorityName="Manton and Oates" authorityYear="1975" box="[1156,1316,1571,1597]" class="Coccolithophyceae" family="Papposphaeraceae" genus="Pappomonas" kingdom="Chromista" order="Coccosphaerales" pageId="2" pageNumber="237" phylum="Haptophyta" rank="species" species="flabellifera">
|
||
<emphasis id="16970607FF8EFFF2FB58DF34FAF9F990" box="[1156,1316,1571,1597]" italics="true" pageId="2" pageNumber="237">P. flabellifera</emphasis>
|
||
</taxonomicName>
|
||
cells examined in preparation for the current paper and also those illustrated in
|
||
<figureCitation id="BCD8C690FF8EFFF2FC14DF78FBE6F92F" box="[968,1083,1640,1666]" captionStart-0="Figs 3–8" captionStart-1="Figs 9–17" captionStartId-0="4.[134,177,1749,1771]" captionStartId-1="8.[132,175,1752,1774]" captionText-0="Figs 3–8. Pappomonas flabellifera TEM whole mounts from the Arctic (NEW). 3 – whole cell with details accounted for in Figs 6–7; the arrow points to the demarcation line between the significantly different sized half blades of the triangle; 4 – whole cell with flagella and a curled up haptonema; 5 – detail from Fig. 4 showing body coccoliths with a fairly extensive central area calcification; reversed printing; 6 – body coccoliths (detail from Fig. 3) showing rim and central area calcification; 7 – oval flagellar pole coccolith with extensive central area calcification; notice the minute elements making up the stalk and the short overlap between stalk and triangle (arrows); 8 – detached armour of coccoliths; circumflagellar coccoliths (arrows) are oval, nearly of the same size as body coccoliths and with fairly extensive calcification." captionText-1="Figs 9–17. Pappomonas flabellifera from Svalbard (9–12), Mexico (13–14) and California (15–17). SEM (9–12) and TEM (13–17) whole mounts. 9 – complete cell showing closely packed body coccoliths and a very well defined ring of circumflagellar coccoliths (ca. 15); 10 – detail of body coccoliths showing the upright position of rim elements; 11 – complete cell; 12 – detail of circumflagellar coccoliths (from Fig. 9) indicating that neighbouring ‘half blades’ of a triangle are joined at a blunt angle; 13 – whole cell very reminiscent of material previously published from S. Africa; see text for further details; 14 – detail of a reduced central appendage formed by two equally sized half blades; 15 – whole cell with flagella and haptonema; 16 – detail from Fig. 15 showing triangles that are reminiscent of those of the S.African type cell; see text for further details; 17 – detail from Fig. 15 of reduced, yet stalked central appendages." pageId="2" pageNumber="237">Figs 3–12</figureCitation>
|
||
come from the North East Water Polynya (
|
||
<collectionCode id="42F242D0FF8EFFF2FC7CDF9BFC39F908" box="[928,996,1675,1701]" country="Australia" httpUri="http://grbio.org/cool/93c9-w2q7" name="University of Newcastle" pageId="2" pageNumber="237">NEW</collectionCode>
|
||
/
|
||
<figureCitation id="BCD8C690FF8EFFF2FC27DF9BFBBFF908" box="[1019,1122,1675,1701]" captionStart="Figs 3–8" captionStartId="4.[134,177,1749,1771]" captionText="Figs 3–8. Pappomonas flabellifera TEM whole mounts from the Arctic (NEW). 3 – whole cell with details accounted for in Figs 6–7; the arrow points to the demarcation line between the significantly different sized half blades of the triangle; 4 – whole cell with flagella and a curled up haptonema; 5 – detail from Fig. 4 showing body coccoliths with a fairly extensive central area calcification; reversed printing; 6 – body coccoliths (detail from Fig. 3) showing rim and central area calcification; 7 – oval flagellar pole coccolith with extensive central area calcification; notice the minute elements making up the stalk and the short overlap between stalk and triangle (arrows); 8 – detached armour of coccoliths; circumflagellar coccoliths (arrows) are oval, nearly of the same size as body coccoliths and with fairly extensive calcification." pageId="2" pageNumber="237">Figs 3–8</figureCitation>
|
||
) and
|
||
<collectingCountry id="5CF49A85FF8EFFF2FB7FDF9BFAD6F908" box="[1187,1291,1675,1701]" name="Svalbard and Jan Mayen" pageId="2" pageNumber="237">Svalbard</collectingCountry>
|
||
(
|
||
<figureCitation id="BCD8C690FF8EFFF2FAC7DF9BFA48F908" box="[1307,1429,1675,1701]" captionStart="Figs 9–17" captionStartId="8.[132,175,1752,1774]" captionText="Figs 9–17. Pappomonas flabellifera from Svalbard (9–12), Mexico (13–14) and California (15–17). SEM (9–12) and TEM (13–17) whole mounts. 9 – complete cell showing closely packed body coccoliths and a very well defined ring of circumflagellar coccoliths (ca. 15); 10 – detail of body coccoliths showing the upright position of rim elements; 11 – complete cell; 12 – detail of circumflagellar coccoliths (from Fig. 9) indicating that neighbouring ‘half blades’ of a triangle are joined at a blunt angle; 13 – whole cell very reminiscent of material previously published from S. Africa; see text for further details; 14 – detail of a reduced central appendage formed by two equally sized half blades; 15 – whole cell with flagella and haptonema; 16 – detail from Fig. 15 showing triangles that are reminiscent of those of the S.African type cell; see text for further details; 17 – detail from Fig. 15 of reduced, yet stalked central appendages." pageId="2" pageNumber="237">Figs 9–12</figureCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph id="245CDA15FF8EFFF4FC8ADFBEFD46FD72" blockId="2.[816,1463,427,1944]" lastBlockId="4.[132,779,223,1567]" lastPageId="4" lastPageNumber="239" pageId="2" pageNumber="237">
|
||
The
|
||
<taxonomicName id="E3E3A196FF8EFFF2FC53DFBEFBF2F965" authorityName="Manton and Oates" authorityYear="1975" box="[911,1071,1710,1736]" class="Coccolithophyceae" family="Papposphaeraceae" genus="Pappomonas" kingdom="Chromista" order="Coccosphaerales" pageId="2" pageNumber="237" phylum="Haptophyta" rank="species" species="flabellifera">
|
||
<emphasis id="16970607FF8EFFF2FC53DFBEFBF2F965" box="[911,1071,1710,1736]" italics="true" pageId="2" pageNumber="237">P. flabellifera</emphasis>
|
||
</taxonomicName>
|
||
body coccolith is oval and dis- plays a fairly narrow variability in length and width in the material examined here (see
|
||
<figureCitation id="BCD8C690FF8EFFF2FB65DFE3FAD3F8A0" box="[1209,1294,1779,1805]" captionStart="Fig" captionStartId="8.[816,851,743,765]" captionTargetBox="[829,1453,240,697]" captionTargetId="figure-476@8.[816,1466,223,712]" captionTargetPageId="8" captionText="Fig. 18. Pappomonas flabellifera (squares) and P. borealis (triangles) body scale dimensions." figureDoi="http://doi.org/10.5281/zenodo.12638354" httpUri="https://zenodo.org/record/12638354/files/figure.png" pageId="2" pageNumber="237">Fig. 18</figureCitation>
|
||
and
|
||
<tableCitation id="6961EFAEFF8EFFF2FA91DFE3FA78F8A0" box="[1357,1445,1779,1805]" captionStart="Table 1" captionStartId="4.[816,872,223,244]" captionText="Table 1. Dimensions of body coccoliths in P. flabellifera and P. borealis." pageId="2" pageNumber="237">Table 1</tableCitation>
|
||
). The rim is more or less perpendicular to the base plate scale (
|
||
<figureCitation id="BCD8C690FF8EFFF2FCA2DE28FC0BF8FF" box="[894,982,1848,1874]" captionStart="Figs 9–17" captionStartId="8.[132,175,1752,1774]" captionText="Figs 9–17. Pappomonas flabellifera from Svalbard (9–12), Mexico (13–14) and California (15–17). SEM (9–12) and TEM (13–17) whole mounts. 9 – complete cell showing closely packed body coccoliths and a very well defined ring of circumflagellar coccoliths (ca. 15); 10 – detail of body coccoliths showing the upright position of rim elements; 11 – complete cell; 12 – detail of circumflagellar coccoliths (from Fig. 9) indicating that neighbouring ‘half blades’ of a triangle are joined at a blunt angle; 13 – whole cell very reminiscent of material previously published from S. Africa; see text for further details; 14 – detail of a reduced central appendage formed by two equally sized half blades; 15 – whole cell with flagella and haptonema; 16 – detail from Fig. 15 showing triangles that are reminiscent of those of the S.African type cell; see text for further details; 17 – detail from Fig. 15 of reduced, yet stalked central appendages." pageId="2" pageNumber="237">Fig. 10</figureCitation>
|
||
) and comprises a distal cycle of 15–20 quasi-pentagonal and closely abutted elements (
|
||
<figureCitation id="BCD8C690FF8EFFF2FABADE4BFA70F8D8" box="[1382,1453,1883,1909]" captionStart="Figs 3–8" captionStartId="4.[134,177,1749,1771]" captionText="Figs 3–8. Pappomonas flabellifera TEM whole mounts from the Arctic (NEW). 3 – whole cell with details accounted for in Figs 6–7; the arrow points to the demarcation line between the significantly different sized half blades of the triangle; 4 – whole cell with flagella and a curled up haptonema; 5 – detail from Fig. 4 showing body coccoliths with a fairly extensive central area calcification; reversed printing; 6 – body coccoliths (detail from Fig. 3) showing rim and central area calcification; 7 – oval flagellar pole coccolith with extensive central area calcification; notice the minute elements making up the stalk and the short overlap between stalk and triangle (arrows); 8 – detached armour of coccoliths; circumflagellar coccoliths (arrows) are oval, nearly of the same size as body coccoliths and with fairly extensive calcification." pageId="2" pageNumber="237">Fig. 6</figureCitation>
|
||
) and interspersed between these a proximal/inner cycle of short rod-like elements (see
|
||
<figureCitation id="BCD8C690FF88FFF4FE22D9CFFDB3FF57" box="[510,622,223,250]" captionStart="Fig" captionStartId="18.[132,167,1724,1746]" captionTargetBox="[157,749,1216,1623]" captionTargetId="figure-8@18.[130,780,1192,1670]" captionTargetPageId="18" captionText="Fig. 58. Interpretation of rim calcification (body coccoliths) in spe- cies of Pappomonas and Papposphaera. A – Pappomonas flabellifera (NEW / micrograph 25036); B – Pappomonas borealis (NEW / micrograph 24879); C – Pappomonas weddellensis (ANT X/3 / mi- crograph 20549); D – Pappomonas garrisonii sp.nov. (ANT X/3 / micrograph 21149); E – Papposphaera sagittifera (NEW / micro- graph 24930); F–G – Balaniger virgulosa HET scale rim (F) and base plate calcification (G) (West Greenland / micrograph 10322)." figureDoi="http://doi.org/10.5281/zenodo.12638364" httpUri="https://zenodo.org/record/12638364/files/figure.png" pageId="4" pageNumber="239">Fig. 58A</figureCitation>
|
||
). The central area calcification comprises rod-shaped elements joined end to end. In larger coccoliths (
|
||
<figureCitation id="BCD8C690FF88FFF4FE2AD835FDE6FE92" box="[502,571,292,319]" captionStart="Figs 3–8" captionStartId="4.[134,177,1749,1771]" captionText="Figs 3–8. Pappomonas flabellifera TEM whole mounts from the Arctic (NEW). 3 – whole cell with details accounted for in Figs 6–7; the arrow points to the demarcation line between the significantly different sized half blades of the triangle; 4 – whole cell with flagella and a curled up haptonema; 5 – detail from Fig. 4 showing body coccoliths with a fairly extensive central area calcification; reversed printing; 6 – body coccoliths (detail from Fig. 3) showing rim and central area calcification; 7 – oval flagellar pole coccolith with extensive central area calcification; notice the minute elements making up the stalk and the short overlap between stalk and triangle (arrows); 8 – detached armour of coccoliths; circumflagellar coccoliths (arrows) are oval, nearly of the same size as body coccoliths and with fairly extensive calcification." pageId="4" pageNumber="239">Fig. 5</figureCitation>
|
||
) there is typically a ring of elements running roughly in parallel with the rim and with a number of more or less parallel lines of elements (3–4) occupying the interior and spanning the entire length of this oval. In smaller coccoliths the central area calcification is limited to 3–4 lines of elements arranged along the longitudinal axis of the coccolith (
|
||
<figureCitation id="BCD8C690FF88FFF4FF51DB07FF39FD9C" box="[141,228,535,561]" captionStart="Figs 9–17" captionStartId="8.[132,175,1752,1774]" captionText="Figs 9–17. Pappomonas flabellifera from Svalbard (9–12), Mexico (13–14) and California (15–17). SEM (9–12) and TEM (13–17) whole mounts. 9 – complete cell showing closely packed body coccoliths and a very well defined ring of circumflagellar coccoliths (ca. 15); 10 – detail of body coccoliths showing the upright position of rim elements; 11 – complete cell; 12 – detail of circumflagellar coccoliths (from Fig. 9) indicating that neighbouring ‘half blades’ of a triangle are joined at a blunt angle; 13 – whole cell very reminiscent of material previously published from S. Africa; see text for further details; 14 – detail of a reduced central appendage formed by two equally sized half blades; 15 – whole cell with flagella and haptonema; 16 – detail from Fig. 15 showing triangles that are reminiscent of those of the S.African type cell; see text for further details; 17 – detail from Fig. 15 of reduced, yet stalked central appendages." pageId="4" pageNumber="239">Fig. 10</figureCitation>
|
||
). In most cases these patterns can be interpret- ed as a ring of elements arranged more or less parallel to the scale rim and with a single line of rods arranged along the longitudinal axis of the scale. Elements from the peripheral ring frequently come into contact with the rim calcification at either end of the coccolith.
|
||
</paragraph>
|
||
<caption id="709C8A9DFF88FFF4FCECD9CFFC40FEBD" pageId="4" pageNumber="239" startId="4.[816,872,223,244]" targetType="table">
|
||
<paragraph id="245CDA15FF88FFF4FCECD9CFFC40FEBD" blockId="4.[816,1461,223,273]" pageId="4" pageNumber="239">
|
||
<emphasis id="16970607FF88FFF4FCECD9CFFC55FF59" bold="true" box="[816,904,223,244]" pageId="4" pageNumber="239">Table 1.</emphasis>
|
||
Dimensions of body coccoliths in
|
||
<taxonomicName id="E3E3A196FF88FFF4FB22D9F0FA58FF59" authorityName="Manton and Oates" authorityYear="1975" box="[1278,1413,223,245]" class="Coccolithophyceae" family="Papposphaeraceae" genus="Pappomonas" kingdom="Chromista" order="Coccosphaerales" pageId="4" pageNumber="239" phylum="Haptophyta" rank="species" species="flabellifera">
|
||
<emphasis id="16970607FF88FFF4FB22D9F0FA58FF59" box="[1278,1413,223,245]" italics="true" pageId="4" pageNumber="239">P. flabellifera</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="E3E3A196FF88FFF4FCECD9ECFC44FEBD" authorityName="Thomsen & Østergaard" authorityYear="2014" baseAuthorityName="Manton, Sutherland and McCully" baseAuthorityYear="1976" box="[816,921,251,273]" class="Coccolithophyceae" family="Papposphaeraceae" genus="Pappomonas" kingdom="Chromista" order="Coccosphaerales" pageId="4" pageNumber="239" phylum="Haptophyta" rank="species" species="borealis">P. borealis</taxonomicName>
|
||
.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="245CDA15FF88FFF4FCECD822FAAEFD88" pageId="4" pageNumber="239">
|
||
<table id="56E328B5FF88000FFCECD822FA50FD8B" box="[816,1421,306,550]" gridcols="4" gridrows="7" pageId="4" pageNumber="239">
|
||
<tr id="9AD3D857FF88000FFCECD822FA50FEE8" box="[816,1421,306,325]" gridrow="0" pageId="4" pageNumber="239" rowspan-1="1">
|
||
<th id="D902B12BFF88000FFCECD822FBDEFEE8" box="[816,1027,306,325]" gridcol="0" gridrow="0" pageId="4" pageNumber="239">Species</th>
|
||
<th id="D902B12BFF88000FFB14D822FAD9FEE8" box="[1224,1284,306,325]" gridcol="2" gridrow="0" pageId="4" pageNumber="239">Length</th>
|
||
<th id="D902B12BFF88000FFA8AD822FA50FEE8" box="[1366,1421,306,325]" gridcol="3" gridrow="0" pageId="4" pageNumber="239">Width</th>
|
||
</tr>
|
||
<tr id="9AD3D857FF88000FFCECD847FA50FEC6" box="[816,1421,343,363]" gridrow="1" pageId="4" pageNumber="239">
|
||
<th id="D902B12BFF88000FFCECD847FBDEFEC6" box="[816,1027,343,363]" gridcol="0" gridrow="1" pageId="4" pageNumber="239">
|
||
<taxonomicName id="E3E3A196FF88FFF4FCECD848FBDEFEC7" authority="Mean" authorityName="Mean" box="[816,1027,343,362]" class="Coccolithophyceae" family="Papposphaeraceae" genus="Pappomonas" kingdom="Chromista" order="Coccosphaerales" pageId="4" pageNumber="239" phylum="Haptophyta" rank="species" species="flabellifera">
|
||
<emphasis id="16970607FF88FFF4FCECD848FBDEFEC7" box="[816,1027,343,362]" italics="true" pageId="4" pageNumber="239">Pappomonas flabellifera</emphasis>
|
||
</taxonomicName>
|
||
</th>
|
||
<td id="D902B12BFF88000FFBC4D847FBA1FEC6" box="[1048,1148,343,363]" gridcol="1" gridrow="1" pageId="4" pageNumber="239">Mean value</td>
|
||
<td id="D902B12BFF88000FFB14D847FAD9FEC6" box="[1224,1284,343,363]" gridcol="2" gridrow="1" pageId="4" pageNumber="239">0.75</td>
|
||
<td id="D902B12BFF88000FFA8AD847FA50FEC6" box="[1366,1421,343,363]" gridcol="3" gridrow="1" pageId="4" pageNumber="239">0.46</td>
|
||
</tr>
|
||
<tr id="9AD3D857FF88000FFCECD86DFA50FE3D" box="[816,1421,381,400]" gridrow="2" pageId="4" pageNumber="239" rowspan-0="1">
|
||
<td id="D902B12BFF88000FFBC4D86DFBA1FE3D" box="[1048,1148,381,400]" gridcol="1" gridrow="2" pageId="4" pageNumber="239">Stand. dev.</td>
|
||
<td id="D902B12BFF88000FFB14D86DFAD9FE3D" box="[1224,1284,381,400]" gridcol="2" gridrow="2" pageId="4" pageNumber="239">0.07</td>
|
||
<td id="D902B12BFF88000FFA8AD86DFA50FE3D" box="[1366,1421,381,400]" gridcol="3" gridrow="2" pageId="4" pageNumber="239">0.05</td>
|
||
</tr>
|
||
<tr id="9AD3D857FF88000FFCECD8B2FA50FE1B" box="[816,1421,418,438]" gridrow="3" pageId="4" pageNumber="239" rowspan-0="1">
|
||
<td id="D902B12BFF88000FFBC4D8B2FBA1FE1B" box="[1048,1148,418,438]" gridcol="1" gridrow="3" pageId="4" pageNumber="239">n</td>
|
||
<td id="D902B12BFF88000FFB14D8B2FAD9FE1B" box="[1224,1284,418,438]" gridcol="2" gridrow="3" pageId="4" pageNumber="239">51</td>
|
||
<td id="D902B12BFF88000FFA8AD8B2FA50FE1B" box="[1366,1421,418,438]" gridcol="3" gridrow="3" pageId="4" pageNumber="239">51</td>
|
||
</tr>
|
||
<tr id="9AD3D857FF88000FFCECD8D8FA50FE76" box="[816,1421,456,475]" gridrow="4" pageId="4" pageNumber="239">
|
||
<th id="D902B12BFF88000FFCECD8D8FBDEFE76" box="[816,1027,456,475]" gridcol="0" gridrow="4" pageId="4" pageNumber="239">
|
||
<taxonomicName id="E3E3A196FF88FFF4FCECD8D8FC37FE76" authority="Mean" authorityName="Thomsen and Ostergaard" authorityYear="2014" baseAuthorityName="Manton, Sutherland and McCully" baseAuthorityYear="1976" box="[816,1002,456,475]" class="Coccolithophyceae" family="Papposphaeraceae" genus="Pappomonas" kingdom="Chromista" order="Coccosphaerales" pageId="4" pageNumber="239" phylum="Haptophyta" rank="species" species="borealis">
|
||
<emphasis id="16970607FF88FFF4FCECD8D8FC37FE76" box="[816,1002,456,475]" italics="true" pageId="4" pageNumber="239">Pappomonas borealis</emphasis>
|
||
</taxonomicName>
|
||
</th>
|
||
<td id="D902B12BFF88000FFBC4D8D8FBA1FE76" box="[1048,1148,456,475]" gridcol="1" gridrow="4" pageId="4" pageNumber="239">Mean value</td>
|
||
<td id="D902B12BFF88000FFB14D8D8FAD9FE76" box="[1224,1284,456,475]" gridcol="2" gridrow="4" pageId="4" pageNumber="239">1.34</td>
|
||
<td id="D902B12BFF88000FFA8AD8D8FA50FE76" box="[1366,1421,456,475]" gridcol="3" gridrow="4" pageId="4" pageNumber="239">0.81</td>
|
||
</tr>
|
||
<tr id="9AD3D857FF88000FFCECD8FDFA50FDAD" box="[816,1421,493,512]" gridrow="5" pageId="4" pageNumber="239" rowspan-0="1">
|
||
<td id="D902B12BFF88000FFBC4D8FDFBA1FDAD" box="[1048,1148,493,512]" gridcol="1" gridrow="5" pageId="4" pageNumber="239">Stand. dev.</td>
|
||
<td id="D902B12BFF88000FFB14D8FDFAD9FDAD" box="[1224,1284,493,512]" gridcol="2" gridrow="5" pageId="4" pageNumber="239">0.15</td>
|
||
<td id="D902B12BFF88000FFA8AD8FDFA50FDAD" box="[1366,1421,493,512]" gridcol="3" gridrow="5" pageId="4" pageNumber="239">0.11</td>
|
||
</tr>
|
||
<tr id="9AD3D857FF88000FFCECDB02FA50FD8B" box="[816,1421,530,550]" gridrow="6" pageId="4" pageNumber="239" rowspan-0="1">
|
||
<td id="D902B12BFF88000FFBC4DB02FBA1FD8B" box="[1048,1148,530,550]" gridcol="1" gridrow="6" pageId="4" pageNumber="239">n</td>
|
||
<td id="D902B12BFF88000FFB14DB02FAD9FD8B" box="[1224,1284,530,550]" gridcol="2" gridrow="6" pageId="4" pageNumber="239">55</td>
|
||
<td id="D902B12BFF88000FFA8ADB02FA50FD8B" box="[1366,1421,530,550]" gridcol="3" gridrow="6" pageId="4" pageNumber="239">55</td>
|
||
</tr>
|
||
</table>
|
||
</paragraph>
|
||
<paragraph id="245CDA15FF88FFF4FF76DBF7FC6BFD48" blockId="4.[132,779,223,1567]" lastBlockId="4.[816,1463,681,1331]" pageId="4" pageNumber="239">
|
||
The circumflagellar coccoliths, typically 10–15, are organized as a fairly compact ring at the anterior cell end (
|
||
<figureCitation id="BCD8C690FF88FFF4FF1EDA3DFECCFCEA" box="[194,273,813,839]" captionStart="Figs 3–8" captionStartId="4.[134,177,1749,1771]" captionText="Figs 3–8. Pappomonas flabellifera TEM whole mounts from the Arctic (NEW). 3 – whole cell with details accounted for in Figs 6–7; the arrow points to the demarcation line between the significantly different sized half blades of the triangle; 4 – whole cell with flagella and a curled up haptonema; 5 – detail from Fig. 4 showing body coccoliths with a fairly extensive central area calcification; reversed printing; 6 – body coccoliths (detail from Fig. 3) showing rim and central area calcification; 7 – oval flagellar pole coccolith with extensive central area calcification; notice the minute elements making up the stalk and the short overlap between stalk and triangle (arrows); 8 – detached armour of coccoliths; circumflagellar coccoliths (arrows) are oval, nearly of the same size as body coccoliths and with fairly extensive calcification." pageId="4" pageNumber="239">Figs 3</figureCitation>
|
||
,
|
||
<figureCitation id="BCD8C690FF88FFF4FEFEDA3DFEBFFCEA" box="[290,354,813,839]" captionStart="Figs 9–17" captionStartId="8.[132,175,1752,1774]" captionText="Figs 9–17. Pappomonas flabellifera from Svalbard (9–12), Mexico (13–14) and California (15–17). SEM (9–12) and TEM (13–17) whole mounts. 9 – complete cell showing closely packed body coccoliths and a very well defined ring of circumflagellar coccoliths (ca. 15); 10 – detail of body coccoliths showing the upright position of rim elements; 11 – complete cell; 12 – detail of circumflagellar coccoliths (from Fig. 9) indicating that neighbouring ‘half blades’ of a triangle are joined at a blunt angle; 13 – whole cell very reminiscent of material previously published from S. Africa; see text for further details; 14 – detail of a reduced central appendage formed by two equally sized half blades; 15 – whole cell with flagella and haptonema; 16 – detail from Fig. 15 showing triangles that are reminiscent of those of the S.African type cell; see text for further details; 17 – detail from Fig. 15 of reduced, yet stalked central appendages." pageId="4" pageNumber="239">9, 11</figureCitation>
|
||
). The hollow process (0.8–10 µm) which is formed by a dense layer of minute rods (
|
||
<figureCitation id="BCD8C690FF88FFF4FD61DA5FFCDCFCC4" box="[701,769,847,873]" captionStart="Figs 3–8" captionStartId="4.[134,177,1749,1771]" captionText="Figs 3–8. Pappomonas flabellifera TEM whole mounts from the Arctic (NEW). 3 – whole cell with details accounted for in Figs 6–7; the arrow points to the demarcation line between the significantly different sized half blades of the triangle; 4 – whole cell with flagella and a curled up haptonema; 5 – detail from Fig. 4 showing body coccoliths with a fairly extensive central area calcification; reversed printing; 6 – body coccoliths (detail from Fig. 3) showing rim and central area calcification; 7 – oval flagellar pole coccolith with extensive central area calcification; notice the minute elements making up the stalk and the short overlap between stalk and triangle (arrows); 8 – detached armour of coccoliths; circumflagellar coccoliths (arrows) are oval, nearly of the same size as body coccoliths and with fairly extensive calcification." pageId="4" pageNumber="239">Fig. 7</figureCitation>
|
||
) is proximally attached to an oval base plate scale (
|
||
<figureCitation id="BCD8C690FF88FFF4FD0ADA62FF02FC02" captionStart="Figs 3–8" captionStartId="4.[134,177,1749,1771]" captionText="Figs 3–8. Pappomonas flabellifera TEM whole mounts from the Arctic (NEW). 3 – whole cell with details accounted for in Figs 6–7; the arrow points to the demarcation line between the significantly different sized half blades of the triangle; 4 – whole cell with flagella and a curled up haptonema; 5 – detail from Fig. 4 showing body coccoliths with a fairly extensive central area calcification; reversed printing; 6 – body coccoliths (detail from Fig. 3) showing rim and central area calcification; 7 – oval flagellar pole coccolith with extensive central area calcification; notice the minute elements making up the stalk and the short overlap between stalk and triangle (arrows); 8 – detached armour of coccoliths; circumflagellar coccoliths (arrows) are oval, nearly of the same size as body coccoliths and with fairly extensive calcification." pageId="4" pageNumber="239">Figs 7 and 8</figureCitation>
|
||
) that does not deviate markedly from an ordinary body coccolith in size, shape or central area calcification. However, the central process is supported also by lines of elements that run more or less perpendicular to the longitudinal axis of the coccolith forming a trans- verse bar. It is also apparent from
|
||
<figureCitation id="BCD8C690FF88FFF4FDC4DD52FD76FBF1" box="[536,683,1090,1117]" captionStart="Figs 3–8" captionStartId="4.[134,177,1749,1771]" captionText="Figs 3–8. Pappomonas flabellifera TEM whole mounts from the Arctic (NEW). 3 – whole cell with details accounted for in Figs 6–7; the arrow points to the demarcation line between the significantly different sized half blades of the triangle; 4 – whole cell with flagella and a curled up haptonema; 5 – detail from Fig. 4 showing body coccoliths with a fairly extensive central area calcification; reversed printing; 6 – body coccoliths (detail from Fig. 3) showing rim and central area calcification; 7 – oval flagellar pole coccolith with extensive central area calcification; notice the minute elements making up the stalk and the short overlap between stalk and triangle (arrows); 8 – detached armour of coccoliths; circumflagellar coccoliths (arrows) are oval, nearly of the same size as body coccoliths and with fairly extensive calcification." pageId="4" pageNumber="239">Figs 7 and 8</figureCitation>
|
||
that the rim in circumflagellar pole coccoliths is less elaborate in comparison with body coccoliths. The pentagonal elements are less abundant and more widely spaced. The central process distally carries a triangular structure formed in most cases by two slightly overlapping, asymmetrically sized and obtusely angled blades (
|
||
<figureCitation id="BCD8C690FF88FFF4FD0ADC02FF3EFAE2" captionStart="Figs 3–8" captionStartId="4.[134,177,1749,1771]" captionText="Figs 3–8. Pappomonas flabellifera TEM whole mounts from the Arctic (NEW). 3 – whole cell with details accounted for in Figs 6–7; the arrow points to the demarcation line between the significantly different sized half blades of the triangle; 4 – whole cell with flagella and a curled up haptonema; 5 – detail from Fig. 4 showing body coccoliths with a fairly extensive central area calcification; reversed printing; 6 – body coccoliths (detail from Fig. 3) showing rim and central area calcification; 7 – oval flagellar pole coccolith with extensive central area calcification; notice the minute elements making up the stalk and the short overlap between stalk and triangle (arrows); 8 – detached armour of coccoliths; circumflagellar coccoliths (arrows) are oval, nearly of the same size as body coccoliths and with fairly extensive calcification." pageId="4" pageNumber="239">Figs 3 and 4</figureCitation>
|
||
) that are terminated distally by a jagged and irregular rim of crystalline facets, and often terminated on either side by a prominent tooth-like projection. The lateral edges of the triangle measure in our material 0.8–1.0 µm and the upper edge from tip to tip 1.0–1.2 µm. The overlap between the central shaft and the two terminal blades is fairly short (
|
||
<figureCitation id="BCD8C690FF88FFF4FE27DF15FD96F9B2" box="[507,587,1541,1567]" captionStart="Figs 3–8" captionStartId="4.[134,177,1749,1771]" captionText="Figs 3–8. Pappomonas flabellifera TEM whole mounts from the Arctic (NEW). 3 – whole cell with details accounted for in Figs 6–7; the arrow points to the demarcation line between the significantly different sized half blades of the triangle; 4 – whole cell with flagella and a curled up haptonema; 5 – detail from Fig. 4 showing body coccoliths with a fairly extensive central area calcification; reversed printing; 6 – body coccoliths (detail from Fig. 3) showing rim and central area calcification; 7 – oval flagellar pole coccolith with extensive central area calcification; notice the minute elements making up the stalk and the short overlap between stalk and triangle (arrows); 8 – detached armour of coccoliths; circumflagellar coccoliths (arrows) are oval, nearly of the same size as body coccoliths and with fairly extensive calcification." pageId="4" pageNumber="239">Figs 7</figureCitation>
|
||
,
|
||
<figureCitation id="BCD8C690FF88FFF4FD87DF15FDA1F9B2" box="[603,636,1541,1567]" captionStart="Figs 9–17" captionStartId="8.[132,175,1752,1774]" captionText="Figs 9–17. Pappomonas flabellifera from Svalbard (9–12), Mexico (13–14) and California (15–17). SEM (9–12) and TEM (13–17) whole mounts. 9 – complete cell showing closely packed body coccoliths and a very well defined ring of circumflagellar coccoliths (ca. 15); 10 – detail of body coccoliths showing the upright position of rim elements; 11 – complete cell; 12 – detail of circumflagellar coccoliths (from Fig. 9) indicating that neighbouring ‘half blades’ of a triangle are joined at a blunt angle; 13 – whole cell very reminiscent of material previously published from S. Africa; see text for further details; 14 – detail of a reduced central appendage formed by two equally sized half blades; 15 – whole cell with flagella and haptonema; 16 – detail from Fig. 15 showing triangles that are reminiscent of those of the S.African type cell; see text for further details; 17 – detail from Fig. 15 of reduced, yet stalked central appendages." pageId="4" pageNumber="239">12</figureCitation>
|
||
). The angle formed by the two blades is at this point approximately 85 degrees.
|
||
</paragraph>
|
||
<paragraph id="245CDA15FF88FFF4FC8ADBFFFADDFA9E" blockId="4.[816,1463,681,1331]" pageId="4" pageNumber="239">
|
||
It is obvious that
|
||
<bibRefCitation id="4072A7E4FF88FFF4FBEADBFFFAAFFCA4" author="Manton I. & Oates K." box="[1078,1394,750,777]" pageId="4" pageNumber="239" pagination="93 - 109" refId="ref9975" refString="Manton I., Oates K. (1975) Fine-structural observations on Papposphaera Tangen from the southern hemisphere and on Pappomonas gen. nov. from South Africa and Greenland. Brit. Phycol. J. 10: 93 - 109" type="journal article" year="1975">Manton and Oates (1975)</bibRefCitation>
|
||
were very cautious when circumscribing
|
||
<taxonomicName id="E3E3A196FF88FFF4FB05DA01FAA5FC86" box="[1241,1400,785,811]" class="Coccolithophyceae" family="Papposphaeraceae" genus="Pappomonas" kingdom="Chromista" order="Coccosphaerales" pageId="4" pageNumber="239" phylum="Haptophyta" rank="species" species="flabellifera">
|
||
<emphasis id="16970607FF88FFF4FB05DA01FAA5FC86" box="[1241,1400,785,811]" italics="true" pageId="4" pageNumber="239">P. flabellifera</emphasis>
|
||
</taxonomicName>
|
||
realizing that it summarizes features sampled across three acknowledged morphotypes. However, it will be evident when dealing with
|
||
<taxonomicName id="E3E3A196FF88FFF4FB91DA69FB17FC3E" authorityName="Thomsen and Ostergaard" authorityYear="2014" baseAuthorityName="Manton, Sutherland and McCully" baseAuthorityYear="1976" box="[1101,1226,889,915]" class="Coccolithophyceae" family="Papposphaeraceae" genus="Pappomonas" kingdom="Chromista" order="Coccosphaerales" pageId="4" pageNumber="239" phylum="Haptophyta" rank="species" species="borealis">
|
||
<emphasis id="16970607FF88FFF4FB91DA69FB17FC3E" box="[1101,1226,889,915]" italics="true" pageId="4" pageNumber="239">P. borealis</emphasis>
|
||
</taxonomicName>
|
||
below that substantial evidence exists in favour of treating
|
||
<taxonomicName id="E3E3A196FF88FFF4FACBDA8CFA48FC18" authorityName="Thomsen and Ostergaard" authorityYear="2014" baseAuthorityName="Manton, Sutherland and McCully" baseAuthorityYear="1976" box="[1303,1429,923,949]" class="Coccolithophyceae" family="Papposphaeraceae" genus="Pappomonas" kingdom="Chromista" order="Coccosphaerales" pageId="4" pageNumber="239" phylum="Haptophyta" rank="species" species="borealis">
|
||
<emphasis id="16970607FF88FFF4FACBDA8CFA48FC18" box="[1303,1429,923,949]" italics="true" pageId="4" pageNumber="239">P. borealis</emphasis>
|
||
</taxonomicName>
|
||
as a separate species and not merely as a variety of
|
||
<emphasis id="16970607FF88FFF4FAAEDAAFFC41FC56" italics="true" pageId="4" pageNumber="239">
|
||
<taxonomicName id="E3E3A196FF88FFF4FAAEDAAFFC4AFC56" class="Coccolithophyceae" family="Papposphaeraceae" genus="Pappomonas" kingdom="Chromista" order="Coccosphaerales" pageId="4" pageNumber="239" phylum="Haptophyta" rank="species" species="flabellifera">P. flabellifera</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
This obviously impacts on the formal circumscription of
|
||
<taxonomicName id="E3E3A196FF88FFF4FC16DD14FBB0FBB0" box="[970,1133,1027,1053]" class="Coccolithophyceae" family="Papposphaeraceae" genus="Pappomonas" kingdom="Chromista" order="Coccosphaerales" pageId="4" pageNumber="239" phylum="Haptophyta" rank="species" species="flabellifera">
|
||
<emphasis id="16970607FF88FFF4FC16DD14FBB0FBB0" box="[970,1133,1027,1053]" italics="true" pageId="4" pageNumber="239">P. flabellifera</emphasis>
|
||
</taxonomicName>
|
||
and calls for an emended description of this taxon. The diagnosis is filtered for
|
||
<taxonomicName id="E3E3A196FF88FFF4FCECDD59FC70FBCE" authorityName="Thomsen and Ostergaard" authorityYear="2014" baseAuthorityName="Manton, Sutherland and McCully" baseAuthorityYear="1976" box="[816,941,1097,1123]" class="Coccolithophyceae" family="Papposphaeraceae" genus="Pappomonas" kingdom="Chromista" order="Coccosphaerales" pageId="4" pageNumber="239" phylum="Haptophyta" rank="species" species="borealis">
|
||
<emphasis id="16970607FF88FFF4FCECDD59FC70FBCE" box="[816,941,1097,1123]" italics="true" pageId="4" pageNumber="239">P. borealis</emphasis>
|
||
</taxonomicName>
|
||
effects in terms of size ranges provided and a lack of informative morphological details caused by the amalgamation of taxonomic units that were in fact too different to be handled at the subspecific level. The emended diagnosis below is building directly upon the original species diagnosis and only revised in places where additional precision is called for.
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |