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<document id="BCA7D4354243C2879797CD44D2DF67C4" ID-DOI="10.1080/00222933.2017.1319520" ID-ISSN="1464-5262" ID-Zenodo-Dep="5181143" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1628687010221" checkinUser="felipe" docAuthor="Jang, Seok Won, Vďačný, Peter, Shazib, Shahed Uddin Ahmed &amp; Shin, Mann Kyoon" docDate="2017" docId="03B0879FFFB1FFA96695FE50FEFB3127" docLanguage="en" docName="JNatHist.51.17-18.939-974.pdf" docOrigin="Journal of Natural History 51 (17 - 18)" docSource="http://dx.doi.org/10.1080/00222933.2017.1319520" docStyle="DocumentStyle:485A7191CD41D6261F41521A3380DC73.9:JNatHist.2017-.journal_article.1cover" docStyleId="485A7191CD41D6261F41521A3380DC73" docStyleName="JNatHist.2017-.journal_article.1cover" docStyleVersion="9" docTitle="Spathidium papilliferum Kahl 1930" docType="treatment" docVersion="3" lastPageNumber="955" masterDocId="FF89FFE7FFBCFFB8672FFF86FFA6375E" masterDocTitle="Linking morphology and molecules: integrative taxonomy of spathidiids (Protista: Ciliophora: Litostomatea) from Korea" masterLastPageNumber="974" masterPageNumber="939" pageNumber="951" updateTime="1699049680283" updateUser="plazi" zenodo-license-document="CLOSED">
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<mods:title id="9332317EAF2173FC2765852D4004A3D2">Linking morphology and molecules: integrative taxonomy of spathidiids (Protista: Ciliophora: Litostomatea) from Korea</mods:title>
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<mods:namePart id="626E14EC3175B4A18202F387CB963A3B">Jang, Seok Won</mods:namePart>
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<mods:namePart id="0BE6E37ACD2F1B8611D3FC528A294268">Shazib, Shahed Uddin Ahmed</mods:namePart>
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<mods:namePart id="09D1B63F9CED156DEC6DAA40B82E54C1">Shin, Mann Kyoon</mods:namePart>
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<paragraph id="8BA63689FFB1FFB56695FE50FCCF36AE" blockId="13.[412,903,470,530]" box="[442,873,470,496]" pageId="13" pageNumber="951">
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<taxonomicName id="4C194D0AFFB1FFB56695FE50FCCF36AE" ID-CoL="6YV86" authority="Kahl, 1930" authorityName="Kahl" authorityYear="1930" box="[442,873,470,496]" class="Gymnostomatea" family="Spathidiidae" genus="Spathidium" kingdom="Chromista" order="Spathidiida" pageId="13" pageNumber="951" phylum="Ciliophora" rank="species" species="papilliferum">
<emphasis id="B96DEA9BFFB1FFB56695FE50FD4336AE" bold="true" box="[442,741,470,496]" italics="true" pageId="13" pageNumber="951">Spathidium papilliferum</emphasis>
<bibRefCitation id="EF884B78FFB1FFB565C1FE51FCCF36AE" author="Kahl A" box="[750,873,471,496]" pageId="13" pageNumber="951" pagination="1 - 180" refId="ref14460" refString="Kahl A. 1930. Urtiere oder Protozoa I: Wimpertiere oder Ciliata (Infusoria) 1. Allgemeiner Teil und Prostomata. Tierwelt Dtl. 18: 1 - 180." type="journal article" year="1930">Kahl, 1930</bibRefCitation>
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<paragraph id="8BA63689FFB1FFB566B3FE7FFC21354C" blockId="13.[412,903,470,530]" box="[412,903,505,530]" pageId="13" pageNumber="951">
(
<figureCitation id="13222A0CFFB1FFB5668BFE7FFDB6354C" box="[420,528,505,530]" captionStart="Figure 5" captionStartId="15.[160,225,1501,1524]" captionText="Figure 5. Spathidium papilliferum, Ssanggyesa temple population from life (ac, g, h) and after protargol impregnation (df, ik). (a) Right side view of a representative individual; (b) resting cyst, 45 µm in diameter; (c) mature and developing extrusomes, about 13 µm long; (d) nuclear apparatus in mid-body; (e, f) variability of body shape and size as well as of nuclear apparatus; (g, h) variability of oral bulge papillae; (i, j) dorsolateral view of ciliary and nuclear pattern in anterior body portion; (k) right side view of ciliary pattern in anterior body portion. CK, circumoral kinety; CV, contractile vacuole; DB, dorsal brush; EX, extrusomes; MA, macronuclear nodules/macronucleus; MI, micronuclei/micronucleus; OB, oral bulge. Scale bars: a, e, f = 100 µm; d, ik = 10 µm." pageId="13" pageNumber="951">Figures 5</figureCitation>
(ak), 6(aj) and 7(ai);
<tableCitation id="C69B0332FFB1FFB56407FE7FFCD9354C" box="[808,895,505,530]" captionStart="Table 2" captionStartId="7.[160,216,142,165]" captionText="Table 2. Morphometric data of Korean populations of Enchelys megaspinata sp. nov. (EM), Spathidium ascendens (SA), S. papilliferum (SPa, population 2), S. polynucleatum comb. nov. (SPo), S. rectitoratum (SR), and Apobryophyllum schmidingeri (AS)." pageId="13" pageNumber="951">Table 2</tableCitation>
)
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<subSubSection id="C3036502FFB1FFB5678FFDABFB2535F0" pageId="13" pageNumber="951" type="reference_group">
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<treatmentCitation id="0AB81098FFB1FFB5678FFDABFDC63518" author="Kahl A" box="[160,608,557,582]" page="164" pageId="13" pageNumber="951" year="1930">
<taxonomicName id="4C194D0AFFB1FFB5678FFDABFDC63518" ID-CoL="6YV86" authority="Kahl, 1930: 164" authorityName="Kahl" authorityPageNumber="164" authorityYear="1930" box="[160,608,557,582]" class="Gymnostomatea" family="Spathidiidae" genus="Spathidium" kingdom="Chromista" order="Spathidiida" pageId="13" pageNumber="951" phylum="Ciliophora" rank="species" species="papilliferum">
<emphasis id="B96DEA9BFFB1FFB5678FFDABFE0E3518" box="[160,424,557,582]" italics="true" pageId="13" pageNumber="951">Spathidium papilliferum</emphasis>
<bibRefCitation id="EF884B78FFB1FFB5669FFDA8FDC63518" author="Kahl A" box="[432,608,558,582]" pageId="13" pageNumber="951" pagination="1 - 180" refId="ref14460" refString="Kahl A. 1930. Urtiere oder Protozoa I: Wimpertiere oder Ciliata (Infusoria) 1. Allgemeiner Teil und Prostomata. Tierwelt Dtl. 18: 1 - 180." type="journal article" year="1930">Kahl, 1930: 164</bibRefCitation>
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, fig. S. 164,
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<paragraph id="8BA63689FFB1FFB5678FFDD6FE6E35D5" blockId="13.[160,1156,557,686]" pageId="13" pageNumber="951">
<treatmentCitationGroup id="AB0911A7FFB1FFB5678FFDD6FE6E35D5" pageId="13" pageNumber="951">
<taxonomicName id="4C194D0AFFB1FFB5678FFDD6FE603537" ID-CoL="6FVRR" authorityName="Foissner" authorityYear="1984" baseAuthorityName="Kahl" baseAuthorityYear="1930" box="[160,454,592,617]" class="Gymnostomatea" family="Spathidiidae" genus="Epispathidium" kingdom="Chromista" order="Spathidiida" pageId="13" pageNumber="951" phylum="Ciliophora" rank="species" species="papilliferum">
<emphasis id="B96DEA9BFFB1FFB5678FFDD6FE603537" box="[160,454,592,617]" italics="true" pageId="13" pageNumber="951">Epispathidium papilliferum</emphasis>
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<treatmentCitation id="0AB81098FFB1FFB566C9FDD7FD123537" author="Foissner W" box="[486,692,593,617]" page="84" pageId="13" pageNumber="951" year="1984">
<bibRefCitation id="EF884B78FFB1FFB566C9FDD7FD123537" author="Foissner W" box="[486,692,593,617]" pageId="13" pageNumber="951" pagination="1 - 165" refId="ref13919" refString="Foissner W. 1984. Infraciliatur, Silberliniensystem und Biometrie einiger neuer und wenig bekannter terrestrischer, limnischer und mariner Ciliaten (Protozoa: Ciliophora) aus den Klassen Kinetofragminophora, Colpodea und Polyhymenophora. Stapfia. 12: 1 - 165." type="journal article" year="1984">Foissner, 1984: 84</bibRefCitation>
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, Abb. 43 (combining author, description of Austrian population)
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<paragraph id="8BA63689FFB1FFB5678FFD13FB2535F0" blockId="13.[160,1156,557,686]" box="[160,1155,661,686]" pageId="13" pageNumber="951">
<treatmentCitationGroup id="AB0911A7FFB1FFB5678FFD13FB2535F0" box="[160,1155,661,686]" pageId="13" pageNumber="951">
<emphasis id="B96DEA9BFFB1FFB5678FFD13FE7D35F0" box="[160,475,661,686]" italics="true" pageId="13" pageNumber="951">
<taxonomicName id="4C194D0AFFB1FFB5678FFD13FE6035F0" ID-CoL="6FVRR" authorityName="Foissner" authorityYear="1984" baseAuthorityName="Kahl" baseAuthorityYear="1930" box="[160,454,661,686]" class="Gymnostomatea" family="Spathidiidae" genus="Epispathidium" kingdom="Chromista" order="Spathidiida" pageId="13" pageNumber="951" phylum="Ciliophora" rank="species" species="papilliferum">Epispathidium papilliferum</taxonomicName>
</emphasis>
<treatmentCitation id="0AB81098FFB1FFB566CAFD10FCE635F0" author="Struder-Kypke MC &amp; Wright A-DG &amp; Foissner W &amp; Chatzinotas A &amp; Lynn DH" box="[485,832,662,686]" page="262" pageId="13" pageNumber="951" year="2006">
<bibRefCitation id="EF884B78FFB1FFB566CAFD10FCE635F0" author="Struder-Kypke MC &amp; Wright A-DG &amp; Foissner W &amp; Chatzinotas A &amp; Lynn DH" box="[485,832,662,686]" pageId="13" pageNumber="951" pagination="261 - 278" refId="ref14947" refString="Struder-Kypke MC, Wright A-DG, Foissner W, Chatzinotas A, Lynn DH. 2006. Molecular phylogeny of litostome ciliates (Ciliophora, Litostomatea) with emphasis on free-living haptorian genera. Protist. 157: 261 - 278." type="journal article" year="2006">Strüder-Kypke et al., 2006: 262</bibRefCitation>
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(18S rRNA gene sequences)
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</paragraph>
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<subSubSection id="C3036502FFB1FFA9678FFD5DFEFB3127" lastPageId="17" lastPageNumber="955" pageId="13" pageNumber="951" type="description">
<paragraph id="8BA63689FFB1FFB5678FFD5DFD9D35AA" blockId="13.[160,1156,731,1379]" box="[160,571,731,756]" pageId="13" pageNumber="951">
<emphasis id="B96DEA9BFFB1FFB5678FFD5DFD9D35AA" bold="true" box="[160,571,731,756]" italics="true" pageId="13" pageNumber="951">Description of Korean populations</emphasis>
</paragraph>
<paragraph id="8BA63689FFB1FFB5678FFD78FC143420" blockId="13.[160,1156,731,1379]" pageId="13" pageNumber="951">
Two populations of this species were studied: one from the Jungjok Mts. (pop. 1) and the other from the surroundings of the Ssanggyesa temple (pop. 2). The populations were very similar; therefore, the description combines all
<emphasis id="B96DEA9BFFB1FFB56403FCC4FCD33405" box="[812,885,834,859]" italics="true" pageId="13" pageNumber="951">in vivo</emphasis>
observations. However, morphometric data were obtained only from the second population.
</paragraph>
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Body size 150190 × 3550 µm, usually near 175 × 45 µm
<emphasis id="B96DEA9BFFB1FFB56477FC0EFC0634FF" box="[856,928,904,929]" italics="true" pageId="13" pageNumber="951">in vivo</emphasis>
and 105185 × 25 65 µm, usually near 140 × 40 µm after protargol impregnation; not contractile but flexible (
<tableCitation id="C69B0332FFB1FFB56788FC48FF5E34B8" box="[167,248,974,998]" captionStart="Table 2" captionStartId="7.[160,216,142,165]" captionText="Table 2. Morphometric data of Korean populations of Enchelys megaspinata sp. nov. (EM), Spathidium ascendens (SA), S. papilliferum (SPa, population 2), S. polynucleatum comb. nov. (SPo), S. rectitoratum (SR), and Apobryophyllum schmidingeri (AS)." pageId="13" pageNumber="951">Table 2</tableCitation>
). Shape narrowly spatulate with a length:width ratio of 3.54.3:
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1
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vivo and 2.7 4.8:1 after protargol impregnation; oral region very conspicuous because oral bulge with two or usually three papillae and about as wide as broadest posterior body portion; neck region rather distinct and flattened laterally; posterior end rounded (
<figureCitation id="13222A0CFFB1FFB564BEFBB0FC533310" box="[913,1013,1078,1102]" captionStart="Figure 5" captionStartId="15.[160,225,1501,1524]" captionText="Figure 5. Spathidium papilliferum, Ssanggyesa temple population from life (ac, g, h) and after protargol impregnation (df, ik). (a) Right side view of a representative individual; (b) resting cyst, 45 µm in diameter; (c) mature and developing extrusomes, about 13 µm long; (d) nuclear apparatus in mid-body; (e, f) variability of body shape and size as well as of nuclear apparatus; (g, h) variability of oral bulge papillae; (i, j) dorsolateral view of ciliary and nuclear pattern in anterior body portion; (k) right side view of ciliary pattern in anterior body portion. CK, circumoral kinety; CV, contractile vacuole; DB, dorsal brush; EX, extrusomes; MA, macronuclear nodules/macronucleus; MI, micronuclei/micronucleus; OB, oral bulge. Scale bars: a, e, f = 100 µm; d, ik = 10 µm." pageId="13" pageNumber="951">Figures 5</figureCitation>
(a,ek), 6(a,b, eh) and 7(a,b,fi)). Nuclear apparatus scattered throughout body, except for the anteriorly flattened cell portion. Number of macronuclear nodules highly variable, ranging from 26 to 79, usually about 50; individual nodules globular, ovate to oblong and about 7.5 × 4.5 µm in size after protargol impregnation; nucleoli small- to medium-sized and rounded. On average 16 micronuclei distributed among macronuclear nodules, globular and about 2.0 µm in diameter after protargol impregnation (
<figureCitation id="13222A0CFFB1FFB56570FA80FD6E3240" box="[607,712,1286,1310]" captionStart="Figure 5" captionStartId="15.[160,225,1501,1524]" captionText="Figure 5. Spathidium papilliferum, Ssanggyesa temple population from life (ac, g, h) and after protargol impregnation (df, ik). (a) Right side view of a representative individual; (b) resting cyst, 45 µm in diameter; (c) mature and developing extrusomes, about 13 µm long; (d) nuclear apparatus in mid-body; (e, f) variability of body shape and size as well as of nuclear apparatus; (g, h) variability of oral bulge papillae; (i, j) dorsolateral view of ciliary and nuclear pattern in anterior body portion; (k) right side view of ciliary pattern in anterior body portion. CK, circumoral kinety; CV, contractile vacuole; DB, dorsal brush; EX, extrusomes; MA, macronuclear nodules/macronucleus; MI, micronuclei/micronucleus; OB, oral bulge. Scale bars: a, e, f = 100 µm; d, ik = 10 µm." pageId="13" pageNumber="951">Figures 5</figureCitation>
(df,i) and 7(ac,eh);
<tableCitation id="C69B0332FFB1FFB56490FA80FBB43240" box="[959,1042,1286,1310]" captionStart="Table 2" captionStartId="7.[160,216,142,165]" captionText="Table 2. Morphometric data of Korean populations of Enchelys megaspinata sp. nov. (EM), Spathidium ascendens (SA), S. papilliferum (SPa, population 2), S. polynucleatum comb. nov. (SPo), S. rectitoratum (SR), and Apobryophyllum schmidingeri (AS)." pageId="13" pageNumber="951">Table 2</tableCitation>
). A single contractile vacuole at posterior body end (
<figureCitation id="13222A0CFFB1FFB565A8FAAFFD48321F" box="[647,750,1321,1345]" captionStart="Figure 5" captionStartId="15.[160,225,1501,1524]" captionText="Figure 5. Spathidium papilliferum, Ssanggyesa temple population from life (ac, g, h) and after protargol impregnation (df, ik). (a) Right side view of a representative individual; (b) resting cyst, 45 µm in diameter; (c) mature and developing extrusomes, about 13 µm long; (d) nuclear apparatus in mid-body; (e, f) variability of body shape and size as well as of nuclear apparatus; (g, h) variability of oral bulge papillae; (i, j) dorsolateral view of ciliary and nuclear pattern in anterior body portion; (k) right side view of ciliary pattern in anterior body portion. CK, circumoral kinety; CV, contractile vacuole; DB, dorsal brush; EX, extrusomes; MA, macronuclear nodules/macronucleus; MI, micronuclei/micronucleus; OB, oral bulge. Scale bars: a, e, f = 100 µm; d, ik = 10 µm." pageId="13" pageNumber="951">Figures 5</figureCitation>
(a,e,f), 6(a,e) and 7(a,b)). Extrusomes densely packed in oral bulge papillae and scattered throughout cytoplasm, impregnate with the protargol method used; oral extrusomes rod-like, almost straight or slightly curved, about 13 µm long
<emphasis id="B96DEA9BFFB3FFB7665CFAC2FE1C3203" box="[371,442,1348,1373]" italics="true" pageId="15" pageNumber="953">in vivo</emphasis>
; developmental cytoplasmic stages either rod-like or composed of a rod-like anterior part and a fusiform or asymmetric posterior part (
<figureCitation id="13222A0CFFB3FFB764E6FAEEFB9732DE" box="[969,1073,1384,1408]" captionStart="Figure 5" captionStartId="15.[160,225,1501,1524]" captionText="Figure 5. Spathidium papilliferum, Ssanggyesa temple population from life (ac, g, h) and after protargol impregnation (df, ik). (a) Right side view of a representative individual; (b) resting cyst, 45 µm in diameter; (c) mature and developing extrusomes, about 13 µm long; (d) nuclear apparatus in mid-body; (e, f) variability of body shape and size as well as of nuclear apparatus; (g, h) variability of oral bulge papillae; (i, j) dorsolateral view of ciliary and nuclear pattern in anterior body portion; (k) right side view of ciliary pattern in anterior body portion. CK, circumoral kinety; CV, contractile vacuole; DB, dorsal brush; EX, extrusomes; MA, macronuclear nodules/macronucleus; MI, micronuclei/micronucleus; OB, oral bulge. Scale bars: a, e, f = 100 µm; d, ik = 10 µm." pageId="15" pageNumber="953">Figures 5</figureCitation>
(c,g,h,j), 6(b,d,fi) and 7(dg,i)). Cortex very flexible, contains several rows of colourless granules between adjacent ciliary rows. Cytoplasm colourless, contains macronuclear nodules, extrusomes, and few to many lipid globules and food vacuoles (
<figureCitation id="13222A0CFFACFFA8647BFA6EFC17315E" box="[852,945,1512,1536]" captionStart="Figure 6" captionStartId="15.[160,225,1126,1149]" captionTargetBox="[162,1152,139,1095]" captionTargetId="figure-207@15.[162,1153,139,1095]" captionTargetPageId="15" captionText="Figure 6. Spathidium papilliferum from life, Jungjok Mts. population (ad) and Ssanggyesa temple population (ej). (a, e) Right side view of representative individuals; (b, fh) detail of anterior body portion showing oral bulge papillae (triangles) studded with extrusomes; (c) dorsal brush bristles; (d, i) extrusomes; (j) resting cyst. CV, contractile vacuole; DB, dorsal brush; EX, extrusomes; LD, lipid droplets. Scale bars: a, e = 100 µm; c = 5 µm; b, d, fj = 10 µm." figureDoi="http://doi.org/10.5281/zenodo.5181149" httpUri="https://zenodo.org/record/5181149/files/figure.png" pageId="16" pageNumber="954">Figure 6</figureCitation>
(a,b,e)). Movement conspicuously slow and worm-like.
</paragraph>
<caption id="DF666601FFB1FFB5678FFA32FEAE31EB" pageId="13" pageNumber="951" startId="13.[160,225,1460,1483]">
<paragraph id="8BA63689FFB1FFB5678FFA32FEAE31EB" blockId="13.[160,1157,1459,1717]" pageId="13" pageNumber="951">
<emphasis id="B96DEA9BFFB1FFB5678FFA32FF5A3295" bold="true" box="[160,252,1460,1483]" pageId="13" pageNumber="951">Figure 4.</emphasis>
<taxonomicName id="4C194D0AFFB1FFB5662AFA35FE7B3294" authorityName="Wenzel" authorityYear="1955" box="[261,477,1459,1482]" class="Gymnostomatea" family="Spathidiidae" genus="Spathidium" kingdom="Chromista" order="Spathidiida" pageId="13" pageNumber="951" phylum="Ciliophora" rank="species" species="ascendens">
<emphasis id="B96DEA9BFFB1FFB5662AFA35FE7B3294" box="[261,477,1459,1482]" italics="true" pageId="13" pageNumber="951">Spathidium ascendens</emphasis>
</taxonomicName>
from life (ah) and after protargol impregnation (im). (a, i) Right side view of representative individuals; (b) detail of posterior body portion showing contractile vacuole; (c) resting cyst; (d, l) detail of anterior body portion showing extrusomes attached to oral bulge; (e) dorsal brush bristles; (f) extrusomes; (g) cortical granulation; (h) epibiotic bacteria; (j, k) right side and dorsolateral views of ciliary pattern in anterior body portion; (m) optical section showing filiform macronucleus and mature rod-like as well as developing fusiform cytoplasmic extrusomes. CK, circumoral kinety; CV, contractile vacuole; DB, dorsal brush; EX, extrusomes; MA, macronucleus; SK, somatic kinety/somatic kineties. Scale bars: a, i = 100 µm; bd, jm = 10 µm; e h = 5 µm.
</paragraph>
</caption>
<caption id="DF666601FFB3FFB7678FFBE0FC9633AD" ID-DOI="http://doi.org/10.5281/zenodo.5181149" ID-Zenodo-Dep="5181149" httpUri="https://zenodo.org/record/5181149/files/figure.png" pageId="15" pageNumber="953" startId="15.[160,225,1126,1149]" targetBox="[162,1152,139,1095]" targetPageId="15">
<paragraph id="8BA63689FFB3FFB7678FFBE0FC9633AD" blockId="15.[160,1156,1126,1267]" pageId="15" pageNumber="953">
<emphasis id="B96DEA9BFFB3FFB7678FFBE0FF5A3323" bold="true" box="[160,252,1126,1149]" pageId="15" pageNumber="953">Figure 6.</emphasis>
<taxonomicName id="4C194D0AFFB3FFB7662AFBE0FE483323" authorityName="Kahl" authorityYear="1930" box="[261,494,1126,1149]" class="Gymnostomatea" family="Spathidiidae" genus="Spathidium" kingdom="Chromista" order="Spathidiida" pageId="15" pageNumber="953" phylum="Ciliophora" rank="species" species="papilliferum">
<emphasis id="B96DEA9BFFB3FFB7662AFBE0FE483323" box="[261,494,1126,1149]" italics="true" pageId="15" pageNumber="953">Spathidium papilliferum</emphasis>
</taxonomicName>
from life, Jungjok Mts. population (ad) and Ssanggyesa temple population (ej). (a, e) Right side view of representative individuals; (b, fh) detail of anterior body portion showing oral bulge papillae (triangles) studded with extrusomes; (c) dorsal brush bristles; (d, i) extrusomes; (j) resting cyst. CV, contractile vacuole; DB, dorsal brush; EX, extrusomes; LD, lipid droplets. Scale bars: a, e = 100 µm; c = 5 µm; b, d, fj = 10 µm.
</paragraph>
</caption>
<caption id="DF666601FFB3FFB7678FFA5BFC1C319C" pageId="15" pageNumber="953" startId="15.[160,225,1501,1524]">
<paragraph id="8BA63689FFB3FFB7678FFA5BFC1C319C" blockId="15.[160,1157,1501,1730]" pageId="15" pageNumber="953">
<emphasis id="B96DEA9BFFB3FFB7678FFA5BFF5932AA" bold="true" box="[160,255,1501,1524]" pageId="15" pageNumber="953">Figure 5.</emphasis>
<taxonomicName id="4C194D0AFFB3FFB76627FA5BFDD032AA" authority=", Ssanggyesa" authorityName="Ssanggyesa" box="[264,630,1501,1524]" class="Gymnostomatea" family="Spathidiidae" genus="Spathidium" kingdom="Chromista" order="Spathidiida" pageId="15" pageNumber="953" phylum="Ciliophora" rank="species" species="papilliferum">
<emphasis id="B96DEA9BFFB3FFB76627FA5BFE5532AA" box="[264,499,1501,1524]" italics="true" pageId="15" pageNumber="953">Spathidium papilliferum</emphasis>
, Ssanggyesa
</taxonomicName>
temple population from life (ac, g, h) and after protargol impregnation (df, ik). (a) Right side view of a representative individual; (b) resting cyst, 45 µm in diameter; (c) mature and developing extrusomes, about 13 µm long; (d) nuclear apparatus in mid-body; (e, f) variability of body shape and size as well as of nuclear apparatus; (g, h) variability of oral bulge papillae; (i, j) dorsolateral view of ciliary and nuclear pattern in anterior body portion; (k) right side view of ciliary pattern in anterior body portion. CK, circumoral kinety; CV, contractile vacuole; DB, dorsal brush; EX, extrusomes; MA, macronuclear nodules/macronucleus; MI, micronuclei/micronucleus; OB, oral bulge. Scale bars: a, e, f = 100 µm; d, ik = 10 µm.
</paragraph>
</caption>
<caption id="DF666601FFACFFA8678FFB12FB8B3226" ID-DOI="http://doi.org/10.5281/zenodo.5181151" ID-Zenodo-Dep="5181151" httpUri="https://zenodo.org/record/5181151/files/figure.png" pageId="16" pageNumber="954" startId="16.[160,225,1172,1195]" targetBox="[183,1153,142,1141]" targetPageId="16">
<paragraph id="8BA63689FFACFFA8678FFB12FB8B3226" blockId="16.[160,1156,1171,1400]" pageId="16" pageNumber="954">
<emphasis id="B96DEA9BFFACFFA8678FFB12FF5C33F5" bold="true" box="[160,250,1172,1195]" pageId="16" pageNumber="954">Figure 7.</emphasis>
<taxonomicName id="4C194D0AFFACFFA8662CFB15FE4E33F4" box="[259,488,1171,1194]" class="Gymnostomatea" family="Spathidiidae" genus="Spathidium" kingdom="Chromista" order="Spathidiida" pageId="16" pageNumber="954" phylum="Ciliophora" rank="species" species="papilliferum">
<emphasis id="B96DEA9BFFACFFA8662CFB15FE4E33F4" box="[259,488,1171,1194]" italics="true" pageId="16" pageNumber="954">Spathidium papilliferum</emphasis>
</taxonomicName>
after protargol impregnation, Ssanggyesa temple population. (a, b) Right side view of representative individuals; (c) nuclear apparatus in mid-body; (d, e) developing and mature cytoplasmic extrusomes; (f) dorsolateral view showing three-rowed dorsal brush and oral bulge papillae studded with extrusomes; (g, h) right and left side views of ciliary pattern in anterior body portion; (i) detail of anterior body portion showing oral bulge papillae with extrusomes. CV, contractile vacuole; CK, circumoral kinety; DB, dorsal brush; EX, extrusomes; MA, macronuclear nodules/macronucleus; MI, micronuclei/micronucleus; SK, somatic kinety/somatic kineties; triangles in (a, b, h, i) denote oral bulge papillae. Scale bars: a, b = 100 µm; ci = 10 µm.
</paragraph>
</caption>
<paragraph id="8BA63689FFACFFA967EFF9ABFC1937B2" blockId="16.[160,1155,1477,1709]" lastBlockId="17.[160,1156,142,444]" lastPageId="17" lastPageNumber="955" pageId="16" pageNumber="954">
Cilia about 8 µm long
<emphasis id="B96DEA9BFFACFFA866EDF9AAFDAD311B" box="[450,523,1580,1605]" italics="true" pageId="16" pageNumber="954">in vivo</emphasis>
, spacing loose to ordinary except for about 10 densely spaced cilia in curved anterior kinety portion. About 26 and 27 equidistant, ordinarily spaced meridional ciliary rows forming an
<taxonomicName id="4C194D0AFFACFFA86554F9F4FCB031D5" authorityName="Foissner" authorityYear="1984" box="[635,790,1650,1675]" class="Gymnostomatea" family="Spathidiidae" genus="Epispathidium" kingdom="Chromista" order="Spathidiida" pageId="16" pageNumber="954" phylum="Ciliophora" rank="genus">
<emphasis id="B96DEA9BFFACFFA86554F9F4FCB031D5" box="[635,790,1650,1675]" italics="true" pageId="16" pageNumber="954">Epispathidium</emphasis>
</taxonomicName>
-like pattern (i.e. anterior portion of right side rows curved dorsally and left side rows curved ventrally to parallel the circumoral kinety;
<figureCitation id="13222A0CFFADFFA96640FF08FE7F37F8" box="[367,473,142,166]" captionStart="Figure 5" captionStartId="15.[160,225,1501,1524]" captionText="Figure 5. Spathidium papilliferum, Ssanggyesa temple population from life (ac, g, h) and after protargol impregnation (df, ik). (a) Right side view of a representative individual; (b) resting cyst, 45 µm in diameter; (c) mature and developing extrusomes, about 13 µm long; (d) nuclear apparatus in mid-body; (e, f) variability of body shape and size as well as of nuclear apparatus; (g, h) variability of oral bulge papillae; (i, j) dorsolateral view of ciliary and nuclear pattern in anterior body portion; (k) right side view of ciliary pattern in anterior body portion. CK, circumoral kinety; CV, contractile vacuole; DB, dorsal brush; EX, extrusomes; MA, macronuclear nodules/macronucleus; MI, micronuclei/micronucleus; OB, oral bulge. Scale bars: a, e, f = 100 µm; d, ik = 10 µm." pageId="17" pageNumber="955">Figures 5</figureCitation>
(i,k) and 7(g,h);
<tableCitation id="C69B0332FFADFFA965ACFF08FD7337F8" box="[643,725,142,166]" captionStart="Table 2" captionStartId="7.[160,216,142,165]" captionText="Table 2. Morphometric data of Korean populations of Enchelys megaspinata sp. nov. (EM), Spathidium ascendens (SA), S. papilliferum (SPa, population 2), S. polynucleatum comb. nov. (SPo), S. rectitoratum (SR), and Apobryophyllum schmidingeri (AS)." pageId="17" pageNumber="955">Table 2</tableCitation>
). Dorsal brush inconspicuous because bristles stump-like and only up to 2.5 µm long
<emphasis id="B96DEA9BFFADFFA965F5FF36FC813797" box="[730,807,176,201]" italics="true" pageId="17" pageNumber="955">in vivo</emphasis>
; dikinetidal; three-rowed; isostichad that is, all rows of similar length (
<figureCitation id="13222A0CFFADFFA965A0FF52FD5D37B2" box="[655,763,212,236]" captionStart="Figure 5" captionStartId="15.[160,225,1501,1524]" captionText="Figure 5. Spathidium papilliferum, Ssanggyesa temple population from life (ac, g, h) and after protargol impregnation (df, ik). (a) Right side view of a representative individual; (b) resting cyst, 45 µm in diameter; (c) mature and developing extrusomes, about 13 µm long; (d) nuclear apparatus in mid-body; (e, f) variability of body shape and size as well as of nuclear apparatus; (g, h) variability of oral bulge papillae; (i, j) dorsolateral view of ciliary and nuclear pattern in anterior body portion; (k) right side view of ciliary pattern in anterior body portion. CK, circumoral kinety; CV, contractile vacuole; DB, dorsal brush; EX, extrusomes; MA, macronuclear nodules/macronucleus; MI, micronuclei/micronucleus; OB, oral bulge. Scale bars: a, e, f = 100 µm; d, ik = 10 µm." pageId="17" pageNumber="955">Figures 5</figureCitation>
(i,j), 6(c) and 7(f)).
</paragraph>
<paragraph id="8BA63689FFADFFA967EFFF70FEE236E2" blockId="17.[160,1156,142,444]" pageId="17" pageNumber="955">
Oral bulge conspicuous because carrying two or usually three papillae; extends over inclined anterior body end (
<figureCitation id="13222A0CFFADFFA96547FE9FFD47366F" box="[616,737,281,305]" captionStart="Figure 5" captionStartId="15.[160,225,1501,1524]" captionText="Figure 5. Spathidium papilliferum, Ssanggyesa temple population from life (ac, g, h) and after protargol impregnation (df, ik). (a) Right side view of a representative individual; (b) resting cyst, 45 µm in diameter; (c) mature and developing extrusomes, about 13 µm long; (d) nuclear apparatus in mid-body; (e, f) variability of body shape and size as well as of nuclear apparatus; (g, h) variability of oral bulge papillae; (i, j) dorsolateral view of ciliary and nuclear pattern in anterior body portion; (k) right side view of ciliary pattern in anterior body portion. CK, circumoral kinety; CV, contractile vacuole; DB, dorsal brush; EX, extrusomes; MA, macronuclear nodules/macronucleus; MI, micronuclei/micronucleus; OB, oral bulge. Scale bars: a, e, f = 100 µm; d, ik = 10 µm." pageId="17" pageNumber="955">Figures 5</figureCitation>
(a,ek), 6(a,b,eh) and 7(a,b,fi)). Circumoral kinety at base of oral bulge; continuous and distinctly separated from ciliary rows; composed of narrowly spaced and perpendicularly oriented dikinetids (
<figureCitation id="13222A0CFFADFFA96787FE07FEB036C7" box="[168,278,385,409]" captionStart="Figure 5" captionStartId="15.[160,225,1501,1524]" captionText="Figure 5. Spathidium papilliferum, Ssanggyesa temple population from life (ac, g, h) and after protargol impregnation (df, ik). (a) Right side view of a representative individual; (b) resting cyst, 45 µm in diameter; (c) mature and developing extrusomes, about 13 µm long; (d) nuclear apparatus in mid-body; (e, f) variability of body shape and size as well as of nuclear apparatus; (g, h) variability of oral bulge papillae; (i, j) dorsolateral view of ciliary and nuclear pattern in anterior body portion; (k) right side view of ciliary pattern in anterior body portion. CK, circumoral kinety; CV, contractile vacuole; DB, dorsal brush; EX, extrusomes; MA, macronuclear nodules/macronucleus; MI, micronuclei/micronucleus; OB, oral bulge. Scale bars: a, e, f = 100 µm; d, ik = 10 µm." pageId="17" pageNumber="955">Figures 5</figureCitation>
(i,k) and 7(g,h)). Nematodesmata not recognisable
<emphasis id="B96DEA9BFFADFFA9644AFE06FC1636C7" box="[869,944,384,409]" italics="true" pageId="17" pageNumber="955">in vivo</emphasis>
or after protargol impregnation.
</paragraph>
<paragraph id="8BA63689FFADFFA9678FFE6EFD72355F" blockId="17.[160,1156,488,686]" box="[160,724,488,513]" pageId="17" pageNumber="955">
<emphasis id="B96DEA9BFFADFFA9678FFE6EFD72355F" bold="true" box="[160,724,488,513]" italics="true" pageId="17" pageNumber="955">Resting cysts of Ssanggyesa temple population</emphasis>
</paragraph>
<paragraph id="8BA63689FFADFFA9678FFD8AFC8D35F0" blockId="17.[160,1156,488,686]" pageId="17" pageNumber="955">
Cysts colourless, about 4045 µm across
<emphasis id="B96DEA9BFFADFFA965ACFD8DFD69357A" box="[643,719,523,548]" italics="true" pageId="17" pageNumber="955">in vivo</emphasis>
; without escape apparatus. Cyst wall inconspicuous because smooth and only about 1 µm thick. Cytoplasm finely granulated, less densely in cyst periphery. Cyst centre studded with globules and macronuclear nodules. Contractile vacuole still present in young cysts. Extrusomes, cortical granules and cilia not recognisable (
<figureCitation id="13222A0CFFADFFA966F9FD10FD9935F0" box="[470,575,662,686]" captionStart="Figure 5" captionStartId="15.[160,225,1501,1524]" captionText="Figure 5. Spathidium papilliferum, Ssanggyesa temple population from life (ac, g, h) and after protargol impregnation (df, ik). (a) Right side view of a representative individual; (b) resting cyst, 45 µm in diameter; (c) mature and developing extrusomes, about 13 µm long; (d) nuclear apparatus in mid-body; (e, f) variability of body shape and size as well as of nuclear apparatus; (g, h) variability of oral bulge papillae; (i, j) dorsolateral view of ciliary and nuclear pattern in anterior body portion; (k) right side view of ciliary pattern in anterior body portion. CK, circumoral kinety; CV, contractile vacuole; DB, dorsal brush; EX, extrusomes; MA, macronuclear nodules/macronucleus; MI, micronuclei/micronucleus; OB, oral bulge. Scale bars: a, e, f = 100 µm; d, ik = 10 µm." pageId="17" pageNumber="955">Figures 5</figureCitation>
(b) and 6(j);
<tableCitation id="C69B0332FFADFFA965E5FD10FCB835F0" box="[714,798,662,686]" captionStart="Table 2" captionStartId="7.[160,216,142,165]" captionText="Table 2. Morphometric data of Korean populations of Enchelys megaspinata sp. nov. (EM), Spathidium ascendens (SA), S. papilliferum (SPa, population 2), S. polynucleatum comb. nov. (SPo), S. rectitoratum (SR), and Apobryophyllum schmidingeri (AS)." pageId="17" pageNumber="955">Table 2</tableCitation>
).
</paragraph>
<paragraph id="8BA63689FFADFFA9678FFD5DFD5B35AA" blockId="17.[160,1156,731,1657]" box="[160,765,731,756]" pageId="17" pageNumber="955">
<emphasis id="B96DEA9BFFADFFA9678FFD5DFD5B35AA" bold="true" box="[160,765,731,756]" italics="true" pageId="17" pageNumber="955">Comparison with original description and remarks</emphasis>
</paragraph>
<paragraph id="8BA63689FFADFFA9678FFD78FEFB3127" blockId="17.[160,1156,731,1657]" pageId="17" pageNumber="955">
Korean populations match Kahl s (1930) description and Foissner s (1984) re-description in their narrowly spatulate body, prominent oral bulge papillae armoured by batteries of rod-shaped extrusomes, several scattered macronuclear nodules, and inconspicuous dorsal brush. However, Korean specimens are smaller than Kahl s individuals (150 190 µm vs 170260 µm), but their size falls well within the range (100200 µm) reported by
<bibRefCitation id="EF884B78FFADFFA967E9FC2DFED2349D" author="Foissner W" box="[198,372,939,963]" pageId="17" pageNumber="955" pagination="1 - 165" refId="ref13919" refString="Foissner W. 1984. Infraciliatur, Silberliniensystem und Biometrie einiger neuer und wenig bekannter terrestrischer, limnischer und mariner Ciliaten (Protozoa: Ciliophora) aus den Klassen Kinetofragminophora, Colpodea und Polyhymenophora. Stapfia. 12: 1 - 165." type="journal article" year="1984">Foissner (1984)</bibRefCitation>
. As concerns the number of macronuclear nodules, there are 710 nodules according to
<bibRefCitation id="EF884B78FFADFFA966BBFC48FDB434B8" author="Kahl A" box="[404,530,974,998]" pageId="17" pageNumber="955" pagination="1 - 180" refId="ref14460" refString="Kahl A. 1930. Urtiere oder Protozoa I: Wimpertiere oder Ciliata (Infusoria) 1. Allgemeiner Teil und Prostomata. Tierwelt Dtl. 18: 1 - 180." type="journal article" year="1930">Kahl (1930)</bibRefCitation>
, while Korean individuals possessed 2679 nodules and Austrian cells about 100 nodules.
<bibRefCitation id="EF884B78FFADFFA96506FC77FD733357" author="Foissner W" box="[553,725,1009,1033]" pageId="17" pageNumber="955" pagination="1 - 165" refId="ref13919" refString="Foissner W. 1984. Infraciliatur, Silberliniensystem und Biometrie einiger neuer und wenig bekannter terrestrischer, limnischer und mariner Ciliaten (Protozoa: Ciliophora) aus den Klassen Kinetofragminophora, Colpodea und Polyhymenophora. Stapfia. 12: 1 - 165." type="journal article" year="1984">Foissner (1984)</bibRefCitation>
assumed that the macronucleus was not fully fragmented in Kahl s specimens, because many minute nodules were scattered among a few larger ones in Austrian cells. On the basis of our observations, we agree with
<bibRefCitation id="EF884B78FFADFFA967F1FBDFFE2A332F" author="Foissner W" box="[222,396,1113,1137]" pageId="17" pageNumber="955" pagination="1 - 165" refId="ref13919" refString="Foissner W. 1984. Infraciliatur, Silberliniensystem und Biometrie einiger neuer und wenig bekannter terrestrischer, limnischer und mariner Ciliaten (Protozoa: Ciliophora) aus den Klassen Kinetofragminophora, Colpodea und Polyhymenophora. Stapfia. 12: 1 - 165." type="journal article" year="1984">Foissner (1984)</bibRefCitation>
that the difference in number of macronuclear nodules can be attributed to life-cycle stage. The length of extrusomes is very similar among all populations: about 13 µm in both Kahl s and Korean specimens and 1012 µm in Foissner s individuals. Likewise, the number of ciliary rows seems to be similar between Korean and
<typeStatus id="54A2882BFFADFFA9678FFB65FF7433A5" box="[160,210,1251,1275]" pageId="17" pageNumber="955">type</typeStatus>
specimens when Kahl s drawing (fig. S. 164,
<emphasis id="B96DEA9BFFADFFA965F0FB64FD4A33A5" box="[735,748,1250,1275]" italics="true" pageId="17" pageNumber="955">7</emphasis>
) is considered. However, Foissner s specimens possessed slightly more ciliary rows than the Korean cells (3236 vs 2627). Our comparison of
<taxonomicName id="4C194D0AFFADFFA96686FAAEFDEC321F" box="[425,586,1320,1345]" class="Gymnostomatea" family="Spathidiidae" genus="Spathidium" kingdom="Chromista" order="Spathidiida" pageId="17" pageNumber="955" phylum="Ciliophora" rank="species" species="papilliferum">
<emphasis id="B96DEA9BFFADFFA96686FAAEFDEC321F" box="[425,586,1320,1345]" italics="true" pageId="17" pageNumber="955">S. papilliferum</emphasis>
</taxonomicName>
populations indicates that this species is either rather variable in body size and number of ciliary rows, or consists of several cryptic species. The second scenario is favoured by molecular analyses (see below). However, the matter is more complex, because Korean populations are not monophyletic and none of them group with the Austrian isolates investigated by
<bibRefCitation id="EF884B78FFADFFA964B3FA35FF4132B0" author="Struder-Kypke MC &amp; Wright A-DG &amp; Foissner W &amp; Chatzinotas A &amp; Lynn DH" pageId="17" pageNumber="955" pagination="261 - 278" refId="ref14947" refString="Struder-Kypke MC, Wright A-DG, Foissner W, Chatzinotas A, Lynn DH. 2006. Molecular phylogeny of litostome ciliates (Ciliophora, Litostomatea) with emphasis on free-living haptorian genera. Protist. 157: 261 - 278." type="journal article" year="2006">Strüder-Kypke et al. (2006)</bibRefCitation>
. It is not possible to solve this taxonomic problem given our present knowledge; this problem is also discussed in the phylogeny section. Because the genus
<taxonomicName id="4C194D0AFFADFFA9678FF99CFE9D316D" authorityName="Foissner" authorityYear="1984" box="[160,315,1562,1587]" class="Gymnostomatea" family="Spathidiidae" genus="Epispathidium" kingdom="Chromista" order="Spathidiida" pageId="17" pageNumber="955" phylum="Ciliophora" rank="genus">
<emphasis id="B96DEA9BFFADFFA9678FF99CFE9D316D" box="[160,315,1562,1587]" italics="true" pageId="17" pageNumber="955">Epispathidium</emphasis>
</taxonomicName>
is polyphyletic, we tentatively prefer Kahl s (1930) original classification of
<taxonomicName id="4C194D0AFFADFFA967ECF9BBFEE33108" box="[195,325,1597,1622]" class="Gymnostomatea" family="Spathidiidae" genus="Spathidium" kingdom="Chromista" order="Spathidiida" pageId="17" pageNumber="955" phylum="Ciliophora" rank="species" species="papilliferum">
<emphasis id="B96DEA9BFFADFFA967ECF9BBFEE33108" box="[195,325,1597,1622]" italics="true" pageId="17" pageNumber="955">papilliferum</emphasis>
</taxonomicName>
in the genus
<taxonomicName id="4C194D0AFFADFFA966DCF9BBFDD63108" authorityName="Dujardin" authorityYear="1841" box="[499,624,1597,1622]" class="Gymnostomatea" family="Spathidiidae" genus="Spathidium" kingdom="Chromista" order="Spathidiida" pageId="17" pageNumber="955" phylum="Ciliophora" rank="genus">
<emphasis id="B96DEA9BFFADFFA966DCF9BBFDD63108" box="[499,624,1597,1622]" italics="true" pageId="17" pageNumber="955">Spathidium</emphasis>
</taxonomicName>
. Further, we consider
<taxonomicName id="4C194D0AFFADFFA96451F9BBFBB83108" box="[894,1054,1597,1622]" class="Gymnostomatea" family="Spathidiidae" genus="Spathidium" kingdom="Chromista" order="Spathidiida" pageId="17" pageNumber="955" phylum="Ciliophora" rank="species" species="papilliferum">
<emphasis id="B96DEA9BFFADFFA96451F9BBFBB83108" box="[894,1054,1597,1622]" italics="true" pageId="17" pageNumber="955">S. papilliferum</emphasis>
</taxonomicName>
to be a collective group.
</paragraph>
</subSubSection>
</treatment>
</document>