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<document id="16C5D61215F31204AA48F7E65F977812" ID-CLB-Dataset="281831" ID-DOI="10.1016/j.jcz.2022.10.008" ID-GBIF-Dataset="cbc8d1b2-efdd-488e-9c9a-04d8e5740627" ID-ISSN="1873-2674" ID-Zenodo-Dep="10375291" 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="1702544581499" checkinUser="felipe" docAuthor="Cepeda, Diego, Gayet, Nicolas, Spedicato, Adriana, Michaud, Emma &amp; Zeppilli, Daniela" docDate="2022" docId="AE098789C674FF802552FD917902FA23" docLanguage="en" docName="ZoolAnz.301.179-195.pdf" docOrigin="Zoologischer Anzeiger 301" docSource="http://dx.doi.org/10.1016/j.jcz.2022.10.008" docStyle="DocumentStyle:32483D6D135DE2FF927BA3F1400E0FFD.1:ZoolAnz.2022-.journal_article" docStyleId="32483D6D135DE2FF927BA3F1400E0FFD" docStyleName="ZoolAnz.2022-.journal_article" docStyleVersion="1" docTitle="Echinoderes angelae Cepeda &amp; Gayet &amp; Spedicato &amp; Michaud &amp; Zeppilli 2022, sp. nov" docType="treatment" docVersion="3" lastPageNumber="186" masterDocId="5230FFF1C675FF872660FFCF783EFFC3" masterDocTitle="Two new species of the Echinoderes coulli-group (Kinorhyncha: Cyclorhagida: Echinoderidae) from a low human-impacted mangrove swamp in French Guiana (western Atlantic Ocean)" masterLastPageNumber="195" masterPageNumber="179" pageNumber="180" updateTime="1702547004206" updateUser="ExternalLinkService">
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<mods:title id="F1DB3CB31F985D2F159534A8D234F88A">Two new species of the Echinoderes coulli-group (Kinorhyncha: Cyclorhagida: Echinoderidae) from a low human-impacted mangrove swamp in French Guiana (western Atlantic Ocean)</mods:title>
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<mods:namePart id="E17162FF1CAA75BCEEA4512F0B8BAD8E">Cepeda, Diego</mods:namePart>
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<mods:namePart id="10C1252D701E88494B11610B4D1ECDFB">Spedicato, Adriana</mods:namePart>
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<mods:namePart id="C221FF041C857157748C0F122CCAA514">Michaud, Emma</mods:namePart>
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<mods:namePart id="098EC6004823D429A29A166C3F1FE559">Zeppilli, Daniela</mods:namePart>
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3.2.
<taxonomicName id="E1A04D1CC674FF862502FD917C28FDB2" ID-CoL="9WJ8W" authority="Cepeda &amp; Gayet &amp; Spedicato &amp; Michaud &amp; Zeppilli, 2022" authorityName="Cepeda &amp; Gayet &amp; Spedicato &amp; Michaud &amp; Zeppilli" authorityYear="2022" box="[866,1046,606,625]" class="Cyclorhagida" family="Echinoderidae" genus="Echinoderes" kingdom="Animalia" order="Echinorhagata" pageId="1" pageNumber="180" phylum="Kinorhyncha" rank="species" species="angelae" status="sp. nov.">Echinoderes angelae</taxonomicName>
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urn:lsid:zoobank.org:act:
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.
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<subSubSection id="6EBA6514C674FF862531FD017C73FD22" box="[849,1101,718,737]" pageId="1" pageNumber="180" type="description">
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(
<figureCitation id="BE9B2A1AC674FF862539FD017B90FD22" box="[857,942,718,737]" captionStart-0="Fig" captionStart-1="Fig" captionStart-2="Fig" captionStart-3="Fig" captionStart-4="Fig" captionStart-5="Fig" captionStart-6="Fig" captionStartId-0="2.[100,130,1889,1906]" captionStartId-1="3.[994,1024,149,166]" captionStartId-2="4.[995,1025,149,166]" captionStartId-3="5.[994,1024,149,166]" captionStartId-4="6.[997,1027,149,166]" captionStartId-5="7.[995,1025,149,166]" captionStartId-6="8.[997,1027,149,166]" captionTargetBox-0="[464,1122,901,1859]" captionTargetBox-1="[101,967,149,1199]" captionTargetBox-2="[102,968,150,977]" captionTargetBox-3="[102,967,150,1427]" captionTargetBox-4="[102,970,150,1260]" captionTargetBox-5="[102,968,150,1247]" captionTargetBox-6="[102,970,150,1291]" captionTargetId-0="figure-514@2.[463,1124,899,1861]" captionTargetId-1="figure-599@3.[100,969,148,1201]" captionTargetId-2="figure-749@4.[100,970,148,979]" captionTargetId-3="figure-287@5.[100,969,148,1429]" captionTargetId-4="figure-385@6.[100,972,148,1261]" captionTargetId-5="figure-378@7.[100,970,148,1249]" captionTargetId-6="figure-503@8.[100,972,148,1293]" captionTargetPageId-0="2" captionTargetPageId-1="3" captionTargetPageId-2="4" captionTargetPageId-3="5" captionTargetPageId-4="6" captionTargetPageId-5="7" captionTargetPageId-6="8" captionText-0="Fig. 2. Diagram of mouth cone, introvert and neck placids in Echinoderes angelae sp. nov. Grey area and thicker line arcs indicate mouth cone and neck placids, respectively. Spinoscalid positions that are inferred by radial symmetry are marked in grey with dashed line." captionText-1="Fig. 3. Line art drawing of Echinoderes angelae sp. nov. A: Ventral female overview; B: dorsal female overview; C: ventral male overview of segments 10–11; D: dorsal male overview of segments 10–11. Abbreviations: dpl, dorsal placid; go, gonopore; lat, lateral accessory tube; lddss, laterodorsal droplet-shaped sensory spot; ldmgco2, laterodorsal modified type 2 glandular cell outlet; ldrss, laterodorsal rounded sensory spot; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; lvgco1, lateroventral type 1 glandular cell outlet; lvmgco2, lateroventral modified type 2 glandular cell outlet; lvt, lateroventral tube; mddss, middorsal droplet-shaped sensory spot; mdgco1, middorsal type 1 glandular cell outlet; mds, middorsal spine; mldss, midlateral droplet-shaped sensory spot; mlmgco2, midlateral modified type 2 glandular cell outlet; ms, muscular scar; mvpl, midventral placid; ppf, primary pectinate fringe; ps, penile spines; S, segment (followed by number of corresponding segment); sddss, subdorsal droplet-shaped sensory spot; sdgco1, subdorsal type 1 glandular cell outlet; sdmgco2, subdorsal modified type 2 glandular cell outlet; sdrss, subdorsal rounded sensory spot; slne, sublateral nephridiopore; sls, sublateral spine; te, tergal extension; tsp, trichoscalid plate; vldss, ventrolateral droplet-shaped sensory spot; vmdss, ventromedial droplet-shaped sensory spot; vmgco1, ventromedial type 1 glandular cell outlet; vmrss, ventromedial rounded sensory spot." captionText-2="Fig. 4. Light micrographs of female holotype MNHN- 655Ma (A, C-D), male paratype MNHN-661Ma (B, E), female paratypes MNHN-656Ma (H) and MNHN-657Ma (F), and additional specimens (G, I) of Echinoderes angelae sp. nov. showing trunk overview, details of mouth cone and neck, epibionts and ingested diatoms. A: Ventral trunk overview; B: mouth cone, showing the outer oral styles; C: dorsal neck view; D: ventral neck view; E: middorsal to laterodorsal view on right side of tergal plate of segments 10–11; F: lateral view of segment 11; G: ingested diatom; H: unknown type of epibionts attached to the posterior edge of segment 11; I: filamentous bacteria attached to the posterior edge of segment 11. Abbreviations: ba, bacteria; bs, basal sheath; dpl, dorsal placid; ep, end-piece; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; mvpl, midventral placid; ps, penile spines; te, tergal extension; tsp, trichoscalid plate." captionText-3="Fig. 5. Light micrographs of female holotype MNHN- 655Ma (E), male paratype MNHN-661Ma (C–D), and additional specimens (A-B, F) of Echinoderes angelae sp. nov. showing trunk cuticular details. A: Middorsal to midlateral view on right side of tergal plate of segments 1–2; B: midlateral to ventromedial view on left side of cuticular plates of segments 5–7; C: ventral view of segments 5–6; D: dorsal view of segments 8–9; E: midlateral to ventromedial view on right side of cuticular plates of segments 9–10 (inset shows the intraspecific variability of the nephridiopore); F: midlateral to ventromedial view on right side of cuticular plates of segments 7–8. Abbreviations: lat, lateral accessory tube; lvt, lateroventral tube; ms, muscular scar; ppf, primary pectinate fringe; slne, sublateral nephridiopore; sls, sublateral spine; numbers after abbreviation indicate corresponding segment; glandular cell outlets are marked with continuous circles, and sensory spots with dashed circles or teardrop-like figures." captionText-4="Fig. 6. Scanning electron micrographs of additional specimens of Echinoderes angelae sp. nov. showing trunk overview and details of introvert, neck, lateral terminal spines and sexually dimorphic features. A: Lateral trunk overview; B: introvert, sector 3; C: introvert, sector 4; D: ventral neck view; E: lateral terminal spine; F: male penile spines; G: female lateral terminal accessory spine. Abbreviations: ba, bacteria; ftz, fringed transverse zone; ltas, lateral terminal accessory spine; lts, lateral terminal spine; mvpl, midventral placid; ne, neck; ps, penile spines; psp, primary spinoscalid; sp, spinoscalid (followed by number of corresponding ring); te, tergal extension; ts, trichoscalid; tsp, trichoscalid plate; lambda marks (̂) indicate attachment point of spinoscalids." captionText-5="Fig. 7. Scanning electron micrographs of additional specimens of Echinoderes angelae sp. nov. showing trunk segments and details of cuticular appendages. A: Subdorsal and laterodorsal sensory spots of segment 1, with the enlarged external ring of micropapillar (inset shows the intraspecific variability of these structures without such kind of external ring); B: ventral view of segments 2–6; C: subdorsal (top) and laterodorsal (bottom) modified type 2 glandular cell outlets of segments 2 and 4 respectively; D: laterodorsal view on left side of tergal plate of segment 2; E: nephridiopore and midlateral dropletshaped sensory spot of segment 9; F: subdorsal to midlateral view on right side of tergal plates of segments 8–9; G: middorsal spine of segment 4; H: subdorsal to midlateral view on left side of tergal plates of segments 3–4; I: laterodorsal tube of segment 10; J: lateral accessory tube of segment 8 (left) and sublateral spine of segment 7 (right); K: subdorsal type 1 glandular cell outlet of segment 9; L: ventral view of segments 10–11; M: subdorsal droplet-shaped sensory spot of segment 9. Abbreviations: go, gonopore; ms, muscular scar; spf, secondary pectinate fringe; modified type 2 glandular cell outlets are marked with continuous circles, and sensory spots with dashed teardrop-like figures." captionText-6="Fig. 8. Scanning electron micrographs showing two types of prokaryotic (likely bacteria) epibionts found on additional specimens of Echinoderes angelae sp. nov. A: Sternal plates of segments 6–7 with filamentous bacteria; B: sternal plates of segments 6–8 with filamentous bacteria; C: filamentous bacteria attached to the ventromedial droplet-shaped sensory spot of segment 6; D: bacteria near the ventrolateral droplet-shaped sensory spot of segment 2; E: bacteria on the muscular scar of segment 6; F: filamentous bacteria attached to the subdorsal enlarged sensory spot of segment 1; G: sternal plates of segments 9–11 and lateral terminal spines with filamentous bacteria; H: bacteria near the lateroventral tube of segment 5; I: bacteria and filamentous bacteria on nephridiopore of segment 9. Abbreviations: tu, tube; arrows indicate the presence and/or attachment point of bacteria and filamentous bacteria respectively." figureDoi-0="http://doi.org/10.5281/zenodo.10375297" figureDoi-1="http://doi.org/10.5281/zenodo.10375299" figureDoi-2="http://doi.org/10.5281/zenodo.10375301" figureDoi-3="http://doi.org/10.5281/zenodo.10375305" figureDoi-4="http://doi.org/10.5281/zenodo.10375307" figureDoi-5="http://doi.org/10.5281/zenodo.10375309" figureDoi-6="http://doi.org/10.5281/zenodo.10375311" httpUri-0="https://zenodo.org/record/10375297/files/figure.png" httpUri-1="https://zenodo.org/record/10375299/files/figure.png" httpUri-2="https://zenodo.org/record/10375301/files/figure.png" httpUri-3="https://zenodo.org/record/10375305/files/figure.png" httpUri-4="https://zenodo.org/record/10375307/files/figure.png" httpUri-5="https://zenodo.org/record/10375309/files/figure.png" httpUri-6="https://zenodo.org/record/10375311/files/figure.png" pageId="1" pageNumber="180">Figs. 2–8</figureCitation>
,
<tableCitation id="6B220324C674FF8625DAFD017C37FD22" box="[954,1033,718,737]" captionStart="Table 1" captionStartId="9.[100,150,150,166]" captionText="Table 1 Morphological measurements and dimensions (in μm) of adult specimens of Echinoderes angelae sp. nov. from the type locality, including number of measured specimens (n), mean value and standard deviation (Sd) of each character, depicted for the holotype and all the measured specimens as a whole. Abbreviations: ac, acicular spine; LA, lateral accessory; LD, laterodorsal; LTAS, lateral terminal accessory spine; LTS, lateral terminal spine; LV, lateroventral; MD, middorsal; MSW, maximum sternal width; S, segment length; SL, sublateral; SW, standard sternal width; t, tube; TL, total trunk length; numbers after abbreviations indicate corresponding segment." pageId="1" pageNumber="180">Tables 1</tableCitation>
and
<tableCitation id="6B220324C674FF862259FD017C7BFD22" box="[1081,1093,718,737]" captionStart="Table 2" captionStartId="9.[100,150,1514,1530]" captionText="Table 2 Summary of nature and arrangement of acicular spines, tubes, sensory spots, glandular cell outlets, nephridiopores and gonopores in adults of Echinoderes angelae sp. nov. Abbreviations: ac, acicular spine; dss, droplet-shaped sensory spot; f, female condition of sexually dimorphic character; gco1, type 1 glandular cell outlet; go, gonopore; LA, lateral accessory; LD, laterodorsal; ltas, lateral terminal accessory spines; lts, lateral terminal spines; LV, lateroventral; m, male condition of sexually dimorphic character; MD, middorsal; mgco2, modified type 2 glandular cell outlet; ML, midlateral; ne, nephridiopore; ps, penile spines; rss, rounded sensory spot; SD, subdorsal; SL, sublateral; t, tube; VL, ventrolateral; VM, ventromedial." pageId="1" pageNumber="180">2</tableCitation>
)
</paragraph>
</subSubSection>
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<emphasis id="14D4EA8DC674FF862552FCCA7C22FCDB" bold="true" box="[818,1052,773,792]" italics="true" pageId="1" pageNumber="180">3.2.1. Material examined</emphasis>
</paragraph>
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<typeStatus id="F91B883DC674FF862531FCEE7B94FCF7" box="[849,938,801,820]" pageId="1" pageNumber="180" type="holotype">Holotype</typeStatus>
, adult female, collected in
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<collectingDate id="425AE9B7C674FF8622CFFCEE7D79FCF7" box="[1199,1351,801,820]" pageId="1" pageNumber="180" value="2017-11">November 2017</collectingDate>
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at the confluence of the
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and Montsin´ery
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22
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27.714
<superScript id="D1D59BD7C674FF86236DFC997D2BFCA0" attach="left" box="[1293,1301,854,867]" fontSize="6" pageId="1" pageNumber="180">′′</superScript>
W
</geoCoordinate>
at the intertidal zone in mud; mounted in
<collectorName id="8B555349C674FF862249FCBA7C87FC4B" box="[1065,1209,885,904]" pageId="1" pageNumber="180">Fluoromount G</collectorName>
®, deposited at
<collectionCode id="40B1AE5AC674FF862330FCBA7DAFFC4B" box="[1360,1425,885,904]" country="France" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34988" lsid="urn:lsid:biocol.org:col:34988" name="Museum National d'Histoire Naturelle" pageId="1" pageNumber="180">MNHN</collectionCode>
under catalogue number: 655
<collectorName id="8B555349C677FF85275FFF5B795EFF64" box="[319,352,148,167]" pageId="2" pageNumber="181">Ma</collectorName>
</materialsCitation>
.
<typeStatus id="F91B883DC677FF852709FF5B79F9FF64" box="[361,455,148,167]" pageId="2" pageNumber="181" type="paratype">Paratypes</typeStatus>
,
<specimenCount id="30A6FD16C677FF8527B2FF5B7A07FF64" box="[466,569,148,167]" count="3" pageId="2" pageNumber="181" type="adult">three adult</specimenCount>
males and
<materialsCitation id="96C83CC2C677FF8524C7FF5B7AFCFF1D" collectingDate="2017-11" collectionCode="MNHN" collectorName="Fluoromount G" country="French Guiana" county="western Atlantic Ocean" latitude="4.897008" location="Cayenne" longLatPrecision="1" longitude="-52.374363" municipality="Rivers" pageId="2" pageNumber="181" specimenCount="5" specimenCount-adult="5" typeStatus="paratype">
<specimenCount id="30A6FD16C677FF8524C7FF5B7B3FFF64" box="[679,769,148,167]" count="5" pageId="2" pageNumber="181" type="adult">five adult</specimenCount>
females, same collecting data as holotype; mounted in
<collectorName id="8B555349C677FF85243EFF7F7AD2FF00" box="[606,748,176,195]" pageId="2" pageNumber="181">Fluoromount G</collectorName>
®, deposited at
<collectionCode id="40B1AE5AC677FF852683FF03791AFF1C" box="[227,292,204,223]" country="France" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34988" lsid="urn:lsid:biocol.org:col:34988" name="Museum National d'Histoire Naturelle" pageId="2" pageNumber="181">MNHN</collectionCode>
under catalogue numbers: 656Ma–663Ma
</materialsCitation>
.
<materialsCitation id="96C83CC2C677FF8524AEFF04789BFE61" collectingDate="2017-11" collectionCode="MNHN" collectorName="Fluoromount G" country="French Guiana" latitude="4.858881" location="Cayenne River" longLatPrecision="1" longitude="-52.399868" municipality="western Atlantic Ocean" pageId="2" pageNumber="181" specimenCount="150" specimenCount-adult="55" specimenCount-exuvia="12" specimenCount-juvenile="83">
Additional material,
<specimenCount id="30A6FD16C677FF85269CFF277975FF39" box="[252,331,231,251]" count="29" pageId="2" pageNumber="181" type="adult">29 adult</specimenCount>
males,
<specimenCount id="30A6FD16C677FF8527F5FF2779DDFF39" box="[405,483,231,251]" count="23" pageId="2" pageNumber="181" type="adult">23 adult</specimenCount>
females,
<specimenCount id="30A6FD16C677FF85245FFF277AB4FF39" box="[575,650,231,251]" count="3" pageId="2" pageNumber="181" type="adult">3 adults</specimenCount>
of unknown sex,
<specimenCount id="30A6FD16C677FF8526EDFECC78C0FED5" box="[141,254,259,278]" count="83" pageId="2" pageNumber="181" type="juvenile">83 juveniles</specimenCount>
and
<specimenCount id="30A6FD16C677FF85274BFECC79B6FED5" box="[299,392,259,278]" count="12" pageId="2" pageNumber="181" type="exuvia">12 exuvia</specimenCount>
, some of them with same collecting data as type material, mounted in
<collectorName id="8B555349C677FF8527F4FED07A17FEF1" box="[404,553,287,306]" pageId="2" pageNumber="181">Fluoromount G</collectorName>
®, others collected in
<date id="521E105FC677FF852604FEF4793EFE8D" box="[100,256,315,334]" pageId="2" pageNumber="181" value="2017-11">
<collectingDate id="425AE9B7C677FF852604FEF4793EFE8D" box="[100,256,315,334]" pageId="2" pageNumber="181" value="2017-11">November 2017</collectingDate>
</date>
ca.
<quantity id="E1589B7AC677FF852750FEF4794EFE8D" box="[304,368,315,334]" metricMagnitude="4" metricUnit="m" metricValue="1.4" pageId="2" pageNumber="181" unit="km" value="14.0">14 km</quantity>
upstream of the
<location id="237F6044C677FF852443FEF47A8DFE8D" LSID="urn:lsid:plazi:treatment:AE098789C674FF802552FD917902FA23:237F6044C677FF852443FEF47A8DFE8D" box="[547,691,315,334]" country="French Guiana" latitude="4.858881" longLatPrecision="1" longitude="-52.399868" municipality="western Atlantic Ocean" name="Cayenne River" pageId="2" pageNumber="181">Cayenne River</location>
,
<collectingCountry id="5EB7760FC677FF8524A1FEF47899FEA9" name="French Guiana" pageId="2" pageNumber="181">French Guiana</collectingCountry>
(
<collectingMunicipality id="C67BACE5C677FF8526D3FE9879AFFEA9" box="[179,401,343,362]" pageId="2" pageNumber="181">western Atlantic Ocean</collectingMunicipality>
):
<geoCoordinate id="43945058C677FF8527C0FE987A72FEA9" box="[416,588,340,362]" degrees="04" direction="north" minutes="51" orientation="latitude" pageId="2" pageNumber="181" precision="1" seconds="31.9716" value="4.858881">
04
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51
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31.9716
<superScript id="D1D59BD7C677FF85244BFE9B7A0DFEA2" attach="left" box="[555,563,340,353]" fontSize="6" pageId="2" pageNumber="181">′′</superScript>
N
</geoCoordinate>
,
<geoCoordinate id="43945058C677FF852434FE987B3CFEA9" box="[596,770,340,362]" degrees="52" direction="west" minutes="23" orientation="longitude" pageId="2" pageNumber="181" precision="1" seconds="59.5248" value="-52.399868">
52
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23
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59.5248
<superScript id="D1D59BD7C677FF852480FE9B7AD6FEA2" attach="left" box="[736,744,340,353]" fontSize="6" pageId="2" pageNumber="181">′′</superScript>
W
</geoCoordinate>
at the intertidal zone in mud; mounted in
<collectorName id="8B555349C677FF852787FEBC7A48FE45" box="[487,630,371,390]" pageId="2" pageNumber="181">Fluoromount G</collectorName>
®, deposited at
<collectionCode id="40B1AE5AC677FF852604FE40789BFE61" box="[100,165,399,418]" country="France" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34988" lsid="urn:lsid:biocol.org:col:34988" name="Museum National d'Histoire Naturelle" pageId="2" pageNumber="181">MNHN</collectionCode>
</materialsCitation>
<materialsCitation id="96C83CC2C677FF8526CEFE4078E8FE35" collectionCode="SEM" country="French Guiana" pageId="2" pageNumber="181" specimenCount="26" specimenCount-adult="18" specimenCount-juvenile="8">
under catalogue numbers: 666Ma–815Ma;
<specimenCount id="30A6FD16C677FF85242FFE407A82FE61" box="[591,700,399,418]" count="7" pageId="2" pageNumber="181" type="adult">seven adult</specimenCount>
males,
<specimenCount id="30A6FD16C677FF852604FE6478F2FE7D" box="[100,204,427,446]" count="8" pageId="2" pageNumber="181" type="adult">eight adult</specimenCount>
females,
<specimenCount id="30A6FD16C677FF85274AFE6479A2FE7D" box="[298,412,427,446]" count="3" pageId="2" pageNumber="181" type="adult">three adults</specimenCount>
of unknown sex and
<specimenCount id="30A6FD16C677FF852411FE647AC0FE7D" box="[625,766,427,446]" count="8" pageId="2" pageNumber="181" type="juvenile">eight juveniles</specimenCount>
, same collecting data as type and remaining additional material, mounted for
<collectionCode id="40B1AE5AC677FF8526C9FE2C78E8FE35" box="[169,214,483,502]" pageId="2" pageNumber="181">SEM</collectionCode>
</materialsCitation>
, deposited at
<collectionCode id="40B1AE5AC677FF852739FE2C7988FE35" box="[345,438,483,502]" country="France" httpUri="http://grbio.org/cool/dcg5-z7hg" name="Institut Francais pour l'Etude de la Mer" pageId="2" pageNumber="181">IFREMER</collectionCode>
.
</paragraph>
</subSubSection>
<caption id="72DF6617C674FF862604F89278F2F862" ID-DOI="http://doi.org/10.5281/zenodo.10375293" ID-Zenodo-Dep="10375293" httpUri="https://zenodo.org/record/10375293/files/figure.png" pageId="1" pageNumber="180" startId="1.[100,130,1885,1902]" targetBox="[359,1228,947,1856]" targetPageId="1" targetType="figure">
<paragraph id="261F369FC674FF862604F89278F2F862" blockId="1.[100,1488,1885,1953]" pageId="1" pageNumber="180">
<emphasis id="14D4EA8DC674FF862604F89278A2F8AD" bold="true" box="[100,156,1885,1902]" pageId="1" pageNumber="180">Fig. 1.</emphasis>
Map showing the sampling location in South America (top left), French Guiana (top right) and the Cayenne Estuary (bottom). Sampling stations are marked red circles; mangrove cover is coloured in dark green. (For interpretation of the references to color in this figure legend, the reader is referred to the Web version of this article.)
</paragraph>
</caption>
<subSubSection id="6EBA6514C677FF852604FDD57D03FF00" pageId="2" pageNumber="181" type="diagnosis">
<paragraph id="261F369FC677FF852604FDD578C5FDEE" blockId="2.[100,771,538,837]" box="[100,251,538,557]" pageId="2" pageNumber="181">
<emphasis id="14D4EA8DC677FF852604FDD578C5FDEE" bold="true" box="[100,251,538,557]" italics="true" pageId="2" pageNumber="181">3.2.2. Diagnosis</emphasis>
</paragraph>
<paragraph id="261F369FC677FF8526E4FDF97D03FF00" blockId="2.[100,771,538,837]" lastBlockId="2.[818,1487,148,195]" pageId="2" pageNumber="181">
<taxonomicName id="E1A04D1CC677FF8526E4FDF978D3FD8A" authorityName="Claparede" authorityYear="1863" box="[132,237,566,585]" class="Cyclorhagida" family="Echinoderidae" genus="Echinoderes" kingdom="Animalia" order="Echinorhagata" pageId="2" pageNumber="181" phylum="Kinorhyncha" rank="genus">
<emphasis id="14D4EA8DC677FF8526E4FDF978D3FD8A" bold="true" box="[132,237,566,585]" italics="true" pageId="2" pageNumber="181">Echinoderes</emphasis>
</taxonomicName>
with short, poorly sclerotized, weakly articulated spines in middorsal position on segment 4 and sublateral position on segments 6–7. Tubes in lateroventral position on segment 5, lateral accessory position on segment 8 and laterodorsal position on segment 10. Modified
<typeStatus id="F91B883DC677FF8526EDFD697888FD7A" box="[141,182,678,697]" pageId="2" pageNumber="181">type</typeStatus>
2 glandular cell outlets in subdorsal position on segments 2 and 5–8, laterodorsal position on segments 2, 4 and 10, midlateral position on segments 2, 6 and 8, and lateroventral position on segments 7–8. Large, rounded sensory spots on segment 1 with a transversal row of conspicuously elongated hairs at the posterior edge of the papillae area. Enlarged, oval sieve plate in sublateral position on segment 9, consisting of an anterior, slightly convex region with numerous pores and a posterior, slightly concave area with a single, central pore.
</paragraph>
</subSubSection>
<subSubSection id="6EBA6514C677FF802552FF287902FA23" lastPageId="7" lastPageNumber="186" pageId="2" pageNumber="181" type="description">
<paragraph id="261F369FC677FF852552FF287BEEFF39" blockId="2.[818,1487,231,307]" box="[818,976,231,250]" pageId="2" pageNumber="181">
<emphasis id="14D4EA8DC677FF852552FF287BEEFF39" bold="true" box="[818,976,231,250]" italics="true" pageId="2" pageNumber="181">3.2.3. Etymology</emphasis>
</paragraph>
<paragraph id="261F369FC677FF852531FECC7D24FEF0" blockId="2.[818,1487,231,307]" pageId="2" pageNumber="181">The species is dedicated to Angela´Cepeda Gomez´, sister of the first author, for all these years of unconditional support.</paragraph>
<paragraph id="261F369FC677FF852552FE987BE9FEA9" blockId="2.[818,1488,343,837]" box="[818,983,343,362]" pageId="2" pageNumber="181">
<emphasis id="14D4EA8DC677FF852552FE987BE9FEA9" bold="true" box="[818,983,343,362]" italics="true" pageId="2" pageNumber="181">3.2.4. Description</emphasis>
</paragraph>
<paragraph id="261F369FC677FF852531FEBC7D67FE7D" blockId="2.[818,1488,343,837]" pageId="2" pageNumber="181">
See
<tableCitation id="6B220324C677FF852516FEBC7B85FE45" box="[886,955,371,390]" captionStart="Table 1" captionStartId="9.[100,150,150,166]" captionText="Table 1 Morphological measurements and dimensions (in μm) of adult specimens of Echinoderes angelae sp. nov. from the type locality, including number of measured specimens (n), mean value and standard deviation (Sd) of each character, depicted for the holotype and all the measured specimens as a whole. Abbreviations: ac, acicular spine; LA, lateral accessory; LD, laterodorsal; LTAS, lateral terminal accessory spine; LTS, lateral terminal spine; LV, lateroventral; MD, middorsal; MSW, maximum sternal width; S, segment length; SL, sublateral; SW, standard sternal width; t, tube; TL, total trunk length; numbers after abbreviations indicate corresponding segment." pageId="2" pageNumber="181">Table 1</tableCitation>
for measurements of selected morphological features and body dimensions, and
<tableCitation id="6B220324C677FF852278FE407C5BFE61" box="[1048,1125,399,418]" captionStart="Table 2" captionStartId="9.[100,150,1514,1530]" captionText="Table 2 Summary of nature and arrangement of acicular spines, tubes, sensory spots, glandular cell outlets, nephridiopores and gonopores in adults of Echinoderes angelae sp. nov. Abbreviations: ac, acicular spine; dss, droplet-shaped sensory spot; f, female condition of sexually dimorphic character; gco1, type 1 glandular cell outlet; go, gonopore; LA, lateral accessory; LD, laterodorsal; ltas, lateral terminal accessory spines; lts, lateral terminal spines; LV, lateroventral; m, male condition of sexually dimorphic character; MD, middorsal; mgco2, modified type 2 glandular cell outlet; ML, midlateral; ne, nephridiopore; ps, penile spines; rss, rounded sensory spot; SD, subdorsal; SL, sublateral; t, tube; VL, ventrolateral; VM, ventromedial." pageId="2" pageNumber="181">Table 2</tableCitation>
for summary of spine, tube, nephridiopore, glandular cell outlet and sensory spot locations.
</paragraph>
<paragraph id="261F369FC677FF852531FE087D5DFD5E" blockId="2.[818,1488,343,837]" pageId="2" pageNumber="181">
Despite the high number of studied specimens, only a few had the head everted, hence only some data of the morphology and appendage arrangement are provided. Ring 00 of mouth cone with nine outer oral styles alternating in size between slightly larger and smaller ones. Outer oral styles composed of two jointed subunits: a rectangular, basal sheath with distal fringe and a triangular, hooked, distally pointed end-piece. Outer oral styles located anterior to each introvert sector, except in the middorsal sector 6 where a style is missing (
<figureCitation id="BE9B2A1AC677FF852299FD457D11FD5E" box="[1273,1327,650,669]" captionStart="Fig" captionStartId="2.[100,130,1889,1906]" captionTargetBox="[464,1122,901,1859]" captionTargetId="figure-514@2.[463,1124,899,1861]" captionTargetPageId="2" captionText="Fig. 2. Diagram of mouth cone, introvert and neck placids in Echinoderes angelae sp. nov. Grey area and thicker line arcs indicate mouth cone and neck placids, respectively. Spinoscalid positions that are inferred by radial symmetry are marked in grey with dashed line." figureDoi="http://doi.org/10.5281/zenodo.10375297" httpUri="https://zenodo.org/record/10375297/files/figure.png" pageId="2" pageNumber="181">Fig. 2</figureCitation>
;
<figureCitation id="BE9B2A1AC677FF85235BFD457D66FD5E" box="[1339,1368,650,669]" captionStart="Fig" captionStartId="4.[995,1025,149,166]" captionTargetBox="[102,968,150,977]" captionTargetId="figure-749@4.[100,970,148,979]" captionTargetPageId="4" captionText="Fig. 4. Light micrographs of female holotype MNHN- 655Ma (A, C-D), male paratype MNHN-661Ma (B, E), female paratypes MNHN-656Ma (H) and MNHN-657Ma (F), and additional specimens (G, I) of Echinoderes angelae sp. nov. showing trunk overview, details of mouth cone and neck, epibionts and ingested diatoms. A: Ventral trunk overview; B: mouth cone, showing the outer oral styles; C: dorsal neck view; D: ventral neck view; E: middorsal to laterodorsal view on right side of tergal plate of segments 10–11; F: lateral view of segment 11; G: ingested diatom; H: unknown type of epibionts attached to the posterior edge of segment 11; I: filamentous bacteria attached to the posterior edge of segment 11. Abbreviations: ba, bacteria; bs, basal sheath; dpl, dorsal placid; ep, end-piece; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; mvpl, midventral placid; ps, penile spines; te, tergal extension; tsp, trichoscalid plate." figureDoi="http://doi.org/10.5281/zenodo.10375301" httpUri="https://zenodo.org/record/10375301/files/figure.png" pageId="2" pageNumber="181">4B</figureCitation>
).
</paragraph>
<paragraph id="261F369FC677FF842531FD697A85F8DE" blockId="2.[818,1488,343,837]" lastBlockId="3.[100,771,1300,1961]" lastPageId="3" lastPageNumber="182" pageId="2" pageNumber="181">
Introvert with seven concentric rings of spinoscalids (one ring of primary spinoscalids and six rings of regular-sized spinoscalids) and 10 longitudinal sectors defined by the arrangement of the primary spinoscalids. Ring 01 with 10 primary spinoscalids, larger than remaining scalids, composed of a basal sheath and a distal, elongated, flexible end-piece. Ring 02 with 10 spinoscalids, arranged as one medially in each sector; spinoscalids on this and following rings are morphologically similar to the primary spinoscalids but smaller. Ring 03 with 20 spinoscalids, arranged as two in each sector. Ring 04 with 10 spinoscalids, arranged as one medially in each sector. Ring 05 with 20 spinoscalids, arranged as two in each sector. Ring 06 with five spinoscalids, arranged as one medially in each odd-numbered sector. Ring 07 with an unknown number of smaller spinoscalids, as only those of sectors 1, 3 and 4 could be observed (sectors 1 and 3 with three spinoscalids, sector 4 with two leaving space for the corresponding trichoscalid and its plate); sectors 9 and 8 are likely similar to sectors 3 and
<quantity id="E1589B7AC676FF8427B7F9C079C4F9E1" box="[471,506,1551,1570]" metricMagnitude="-1" metricUnit="m" metricValue="1.016" pageId="3" pageNumber="182" unit="in" value="4.0">4 in</quantity>
this arrangement due to the typical radial symmetry of the kinorhynch introvert. Scalid arrangement identical in all sectors from rings 02–05, with one unpaired scalid in ring 02 followed by a quincunx in rings 03–05; in sector 1 (and likely 9), ring 06 with an unpaired scalid followed by three smaller scalids in ring 07; in sector 4 (and likely 8), ring 06 without scalids followed by two smaller scalids in ring 07 leaving space for the corresponding trichoscalid and its plate; rings 06–07 unknown for remaining sectors. A fringed ring around the introvert at the level of rings 04–05, with tips more elongated in sectors where a spinoscalid is missing in ring 06 (
<figureCitation id="BE9B2A1AC676FF842459F8C57A51F8DE" box="[569,623,1802,1821]" captionStart="Fig" captionStartId="2.[100,130,1889,1906]" captionTargetBox="[464,1122,901,1859]" captionTargetId="figure-514@2.[463,1124,899,1861]" captionTargetPageId="2" captionText="Fig. 2. Diagram of mouth cone, introvert and neck placids in Echinoderes angelae sp. nov. Grey area and thicker line arcs indicate mouth cone and neck placids, respectively. Spinoscalid positions that are inferred by radial symmetry are marked in grey with dashed line." figureDoi="http://doi.org/10.5281/zenodo.10375297" httpUri="https://zenodo.org/record/10375297/files/figure.png" pageId="3" pageNumber="182">Fig. 2</figureCitation>
;
<figureCitation id="BE9B2A1AC676FF84241BF8C57A8EF8DE" box="[635,688,1802,1821]" captionStart="Fig" captionStartId="6.[997,1027,149,166]" captionTargetBox="[102,970,150,1260]" captionTargetId="figure-385@6.[100,972,148,1261]" captionTargetPageId="6" captionText="Fig. 6. Scanning electron micrographs of additional specimens of Echinoderes angelae sp. nov. showing trunk overview and details of introvert, neck, lateral terminal spines and sexually dimorphic features. A: Lateral trunk overview; B: introvert, sector 3; C: introvert, sector 4; D: ventral neck view; E: lateral terminal spine; F: male penile spines; G: female lateral terminal accessory spine. Abbreviations: ba, bacteria; ftz, fringed transverse zone; ltas, lateral terminal accessory spine; lts, lateral terminal spine; mvpl, midventral placid; ne, neck; ps, penile spines; psp, primary spinoscalid; sp, spinoscalid (followed by number of corresponding ring); te, tergal extension; ts, trichoscalid; tsp, trichoscalid plate; lambda marks (̂) indicate attachment point of spinoscalids." figureDoi="http://doi.org/10.5281/zenodo.10375307" httpUri="https://zenodo.org/record/10375307/files/figure.png" pageId="3" pageNumber="182">6B–C</figureCitation>
).
</paragraph>
<caption id="72DF6617C677FF852604F8AE7C32F84F" ID-DOI="http://doi.org/10.5281/zenodo.10375297" ID-Zenodo-Dep="10375297" httpUri="https://zenodo.org/record/10375297/files/figure.png" pageId="2" pageNumber="181" startId="2.[100,130,1889,1906]" targetBox="[464,1122,901,1859]" targetPageId="2" targetType="figure">
<paragraph id="261F369FC677FF852604F8AE7C32F84F" blockId="2.[100,1488,1889,1932]" pageId="2" pageNumber="181">
<emphasis id="14D4EA8DC677FF852604F8AE78A1F8B1" bold="true" box="[100,159,1889,1906]" pageId="2" pageNumber="181">Fig. 2.</emphasis>
Diagram of mouth cone, introvert and neck placids in
<taxonomicName id="E1A04D1CC677FF85241FF8AE7B1FF8B1" authority="Cepeda &amp; Gayet &amp; Spedicato &amp; Michaud &amp; Zeppilli, 2022" authorityName="Cepeda &amp; Gayet &amp; Spedicato &amp; Michaud &amp; Zeppilli" authorityYear="2022" box="[639,801,1889,1906]" class="Cyclorhagida" family="Echinoderidae" genus="Echinoderes" kingdom="Animalia" order="Echinorhagata" pageId="2" pageNumber="181" phylum="Kinorhyncha" rank="species" species="angelae" status="sp. nov.">
<emphasis id="14D4EA8DC677FF85241FF8AE7B1FF8B1" bold="true" box="[639,801,1889,1906]" italics="true" pageId="2" pageNumber="181">Echinoderes angelae</emphasis>
</taxonomicName>
<taxonomicNameLabel id="0FE757F6C677FF852548F8AE7B52F8B2" box="[808,876,1889,1905]" pageId="2" pageNumber="181" rank="species">sp. nov.</taxonomicNameLabel>
Grey area and thicker line arcs indicate mouth cone and neck placids, respectively. Spinoscalid positions that are inferred by radial symmetry are marked in grey with dashed line.
</paragraph>
</caption>
<caption id="72DF6617C676FF842582FF5A7D2BFCFE" ID-DOI="http://doi.org/10.5281/zenodo.10375299" ID-Zenodo-Dep="10375299" httpUri="https://zenodo.org/record/10375299/files/figure.png" pageId="3" pageNumber="182" startId="3.[994,1024,149,166]" targetBox="[101,967,149,1199]" targetPageId="3" targetType="figure">
<paragraph id="261F369FC676FF842582FF5A7D2BFCFE" blockId="3.[994,1489,149,830]" pageId="3" pageNumber="182">
<emphasis id="14D4EA8DC676FF842582FF5A7C25FF65" bold="true" box="[994,1051,149,166]" pageId="3" pageNumber="182">Fig. 3.</emphasis>
Line art drawing of
<taxonomicName id="E1A04D1CC676FF8422AEFF5A7D51FF65" authority="Cepeda &amp; Gayet &amp; Spedicato &amp; Michaud &amp; Zeppilli, 2022" authorityName="Cepeda &amp; Gayet &amp; Spedicato &amp; Michaud &amp; Zeppilli" authorityYear="2022" box="[1230,1391,149,166]" class="Cyclorhagida" family="Echinoderidae" genus="Echinoderes" kingdom="Animalia" order="Echinorhagata" pageId="3" pageNumber="182" phylum="Kinorhyncha" rank="species" species="angelae" status="sp. nov.">
<emphasis id="14D4EA8DC676FF8422AEFF5A7D51FF65" bold="true" box="[1230,1391,149,166]" italics="true" pageId="3" pageNumber="182">Echinoderes angelae</emphasis>
</taxonomicName>
<taxonomicNameLabel id="0FE757F6C676FF842315FF597D86FF65" box="[1397,1464,150,166]" pageId="3" pageNumber="182" rank="species">sp. nov.</taxonomicNameLabel>
A: Ventral female overview; B: dorsal female overview; C: ventral male overview of segments 10–11; D: dorsal male overview of segments 10–11. Abbreviations: dpl, dorsal placid; go, gonopore; lat, lateral accessory tube; lddss, laterodorsal droplet-shaped sensory spot; ldmgco2, laterodorsal modified type 2 glandular cell outlet; ldrss, laterodorsal rounded sensory spot; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; lvgco1, lateroventral type 1 glandular cell outlet; lvmgco2, lateroventral modified type 2 glandular cell outlet; lvt, lateroventral tube; mddss, middorsal droplet-shaped sensory spot; mdgco1, middorsal type 1 glandular cell outlet; mds, middorsal spine; mldss, midlateral droplet-shaped sensory spot; mlmgco2, midlateral modified type 2 glandular cell outlet; ms, muscular scar; mvpl, midventral placid; ppf, primary pectinate fringe; ps, penile spines; S, segment (followed by number of corresponding segment); sddss, subdorsal droplet-shaped sensory spot; sdgco1, subdorsal type 1 glandular cell outlet; sdmgco2, subdorsal modified type 2 glandular cell outlet; sdrss, subdorsal rounded sensory spot; slne, sublateral nephridiopore; sls, sublateral spine; te, tergal extension; tsp, trichoscalid plate; vldss, ventrolateral droplet-shaped sensory spot; vmdss, ventromedial droplet-shaped sensory spot; vmgco1, ventromedial type 1 glandular cell outlet; vmrss, ventromedial rounded sensory spot.
</paragraph>
</caption>
<paragraph id="261F369FC676FF8426E4F8E97B71FA80" blockId="3.[100,771,1300,1961]" lastBlockId="3.[818,1488,1300,1961]" pageId="3" pageNumber="182">
Neck with 16 trapezoidal placids, wider at the base, closely adjacent each other, with distinct joint between the neck and first trunk segment. Midventral placid widest (ca. 10–11 μm at base), remaining ones slightly narrower (ca. 6–8 μm at base). A ring of six short, superficially haired trichoscalids associated with the placids, attached to quadrangular trichoscalid plates in sectors 2, 4, 5, 7, 8 and 10 (
<figureCitation id="BE9B2A1AC676FF84229BFADB7D11FAE4" box="[1275,1327,1300,1319]" captionStart="Fig" captionStartId="2.[100,130,1889,1906]" captionTargetBox="[464,1122,901,1859]" captionTargetId="figure-514@2.[463,1124,899,1861]" captionTargetPageId="2" captionText="Fig. 2. Diagram of mouth cone, introvert and neck placids in Echinoderes angelae sp. nov. Grey area and thicker line arcs indicate mouth cone and neck placids, respectively. Spinoscalid positions that are inferred by radial symmetry are marked in grey with dashed line." figureDoi="http://doi.org/10.5281/zenodo.10375297" httpUri="https://zenodo.org/record/10375297/files/figure.png" pageId="3" pageNumber="182">Fig. 2</figureCitation>
;
<figureCitation id="BE9B2A1AC676FF84235BFADB7D7AFAE4" box="[1339,1348,1300,1319]" captionStart="Fig" captionStartId="3.[994,1024,149,166]" captionTargetBox="[101,967,149,1199]" captionTargetId="figure-599@3.[100,969,148,1201]" captionTargetPageId="3" captionText="Fig. 3. Line art drawing of Echinoderes angelae sp. nov. A: Ventral female overview; B: dorsal female overview; C: ventral male overview of segments 10–11; D: dorsal male overview of segments 10–11. Abbreviations: dpl, dorsal placid; go, gonopore; lat, lateral accessory tube; lddss, laterodorsal droplet-shaped sensory spot; ldmgco2, laterodorsal modified type 2 glandular cell outlet; ldrss, laterodorsal rounded sensory spot; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; lvgco1, lateroventral type 1 glandular cell outlet; lvmgco2, lateroventral modified type 2 glandular cell outlet; lvt, lateroventral tube; mddss, middorsal droplet-shaped sensory spot; mdgco1, middorsal type 1 glandular cell outlet; mds, middorsal spine; mldss, midlateral droplet-shaped sensory spot; mlmgco2, midlateral modified type 2 glandular cell outlet; ms, muscular scar; mvpl, midventral placid; ppf, primary pectinate fringe; ps, penile spines; S, segment (followed by number of corresponding segment); sddss, subdorsal droplet-shaped sensory spot; sdgco1, subdorsal type 1 glandular cell outlet; sdmgco2, subdorsal modified type 2 glandular cell outlet; sdrss, subdorsal rounded sensory spot; slne, sublateral nephridiopore; sls, sublateral spine; te, tergal extension; tsp, trichoscalid plate; vldss, ventrolateral droplet-shaped sensory spot; vmdss, ventromedial droplet-shaped sensory spot; vmgco1, ventromedial type 1 glandular cell outlet; vmrss, ventromedial rounded sensory spot." figureDoi="http://doi.org/10.5281/zenodo.10375299" httpUri="https://zenodo.org/record/10375299/files/figure.png" pageId="3" pageNumber="182">3</figureCitation>
A-B; 4C-D; 6A- D).
</paragraph>
<paragraph id="261F369FC676FF832531FA837B3DFB5B" blockId="3.[818,1488,1300,1961]" lastBlockId="4.[100,771,1073,1957]" lastPageId="4" lastPageNumber="183" pageId="3" pageNumber="182">
Trunk fusiform, heart-shaped in cross-section, composed of 11 segments. Segments 1–2 closed, ring-like cuticular plates; remaining ones with one tergal and two sternal cuticular plates (
<figureCitation id="BE9B2A1AC676FF842369FA4B7D54FA54" box="[1289,1386,1412,1431]" captionStart="Fig" captionStartId="3.[994,1024,149,166]" captionTargetBox="[101,967,149,1199]" captionTargetId="figure-599@3.[100,969,148,1201]" captionTargetPageId="3" captionText="Fig. 3. Line art drawing of Echinoderes angelae sp. nov. A: Ventral female overview; B: dorsal female overview; C: ventral male overview of segments 10–11; D: dorsal male overview of segments 10–11. Abbreviations: dpl, dorsal placid; go, gonopore; lat, lateral accessory tube; lddss, laterodorsal droplet-shaped sensory spot; ldmgco2, laterodorsal modified type 2 glandular cell outlet; ldrss, laterodorsal rounded sensory spot; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; lvgco1, lateroventral type 1 glandular cell outlet; lvmgco2, lateroventral modified type 2 glandular cell outlet; lvt, lateroventral tube; mddss, middorsal droplet-shaped sensory spot; mdgco1, middorsal type 1 glandular cell outlet; mds, middorsal spine; mldss, midlateral droplet-shaped sensory spot; mlmgco2, midlateral modified type 2 glandular cell outlet; ms, muscular scar; mvpl, midventral placid; ppf, primary pectinate fringe; ps, penile spines; S, segment (followed by number of corresponding segment); sddss, subdorsal droplet-shaped sensory spot; sdgco1, subdorsal type 1 glandular cell outlet; sdmgco2, subdorsal modified type 2 glandular cell outlet; sdrss, subdorsal rounded sensory spot; slne, sublateral nephridiopore; sls, sublateral spine; te, tergal extension; tsp, trichoscalid plate; vldss, ventrolateral droplet-shaped sensory spot; vmdss, ventromedial droplet-shaped sensory spot; vmgco1, ventromedial type 1 glandular cell outlet; vmrss, ventromedial rounded sensory spot." figureDoi="http://doi.org/10.5281/zenodo.10375299" httpUri="https://zenodo.org/record/10375299/files/figure.png" pageId="3" pageNumber="182">Fig. 3A–D</figureCitation>
;
<figureCitation id="BE9B2A1AC676FF842315FA4B7DAFFA54" box="[1397,1425,1412,1431]" captionStart="Fig" captionStartId="4.[995,1025,149,166]" captionTargetBox="[102,968,150,977]" captionTargetId="figure-749@4.[100,970,148,979]" captionTargetPageId="4" captionText="Fig. 4. Light micrographs of female holotype MNHN- 655Ma (A, C-D), male paratype MNHN-661Ma (B, E), female paratypes MNHN-656Ma (H) and MNHN-657Ma (F), and additional specimens (G, I) of Echinoderes angelae sp. nov. showing trunk overview, details of mouth cone and neck, epibionts and ingested diatoms. A: Ventral trunk overview; B: mouth cone, showing the outer oral styles; C: dorsal neck view; D: ventral neck view; E: middorsal to laterodorsal view on right side of tergal plate of segments 10–11; F: lateral view of segment 11; G: ingested diatom; H: unknown type of epibionts attached to the posterior edge of segment 11; I: filamentous bacteria attached to the posterior edge of segment 11. Abbreviations: ba, bacteria; bs, basal sheath; dpl, dorsal placid; ep, end-piece; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; mvpl, midventral placid; ps, penile spines; te, tergal extension; tsp, trichoscalid plate." figureDoi="http://doi.org/10.5281/zenodo.10375301" httpUri="https://zenodo.org/record/10375301/files/figure.png" pageId="3" pageNumber="182">4A</figureCitation>
;
<figureCitation id="BE9B2A1AC676FF8423FDFA4C7D99FA55" box="[1437,1447,1411,1430]" captionStart="Fig" captionStartId="5.[994,1024,149,166]" captionTargetBox="[102,967,150,1427]" captionTargetId="figure-287@5.[100,969,148,1429]" captionTargetPageId="5" captionText="Fig. 5. Light micrographs of female holotype MNHN- 655Ma (E), male paratype MNHN-661Ma (C–D), and additional specimens (A-B, F) of Echinoderes angelae sp. nov. showing trunk cuticular details. A: Middorsal to midlateral view on right side of tergal plate of segments 1–2; B: midlateral to ventromedial view on left side of cuticular plates of segments 5–7; C: ventral view of segments 5–6; D: dorsal view of segments 8–9; E: midlateral to ventromedial view on right side of cuticular plates of segments 9–10 (inset shows the intraspecific variability of the nephridiopore); F: midlateral to ventromedial view on right side of cuticular plates of segments 7–8. Abbreviations: lat, lateral accessory tube; lvt, lateroventral tube; ms, muscular scar; ppf, primary pectinate fringe; slne, sublateral nephridiopore; sls, sublateral spine; numbers after abbreviation indicate corresponding segment; glandular cell outlets are marked with continuous circles, and sensory spots with dashed circles or teardrop-like figures." figureDoi="http://doi.org/10.5281/zenodo.10375305" httpUri="https://zenodo.org/record/10375305/files/figure.png" pageId="3" pageNumber="182">5</figureCitation>
A-F; 6A; 7B). Maximum sternal width at segment 8, relatively narrow compared to total trunk length (MSW8/TL average ratio = 19.6%). Cuticular hairs throughout segments 2–11 (absent on segment 1), acicular, non-bracteate, emerging from rounded to slightly oval perforation sites. Cuticular hairs arranged as 6–12 transversal, uninterrupted rows covering the cuticular surface of segments 2–3 (except in the ventromedial region where hairs are missing); as 6–16 transversal rows that become wavy at subdorsal and ventrolateral to ventromedial regions, interrupted by large, oval, muscular scars in laterodorsal position throughout segments 4–10; as 6–8 transversal, uninterrupted rows covering the cuticular surface of segment 11 (hairs of this segment conspicuously shorter) (
<figureCitation id="BE9B2A1AC676FF84227EF91D7CBCF925" box="[1054,1154,1746,1766]" captionStart="Fig" captionStartId="3.[994,1024,149,166]" captionTargetBox="[101,967,149,1199]" captionTargetId="figure-599@3.[100,969,148,1201]" captionTargetPageId="3" captionText="Fig. 3. Line art drawing of Echinoderes angelae sp. nov. A: Ventral female overview; B: dorsal female overview; C: ventral male overview of segments 10–11; D: dorsal male overview of segments 10–11. Abbreviations: dpl, dorsal placid; go, gonopore; lat, lateral accessory tube; lddss, laterodorsal droplet-shaped sensory spot; ldmgco2, laterodorsal modified type 2 glandular cell outlet; ldrss, laterodorsal rounded sensory spot; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; lvgco1, lateroventral type 1 glandular cell outlet; lvmgco2, lateroventral modified type 2 glandular cell outlet; lvt, lateroventral tube; mddss, middorsal droplet-shaped sensory spot; mdgco1, middorsal type 1 glandular cell outlet; mds, middorsal spine; mldss, midlateral droplet-shaped sensory spot; mlmgco2, midlateral modified type 2 glandular cell outlet; ms, muscular scar; mvpl, midventral placid; ppf, primary pectinate fringe; ps, penile spines; S, segment (followed by number of corresponding segment); sddss, subdorsal droplet-shaped sensory spot; sdgco1, subdorsal type 1 glandular cell outlet; sdmgco2, subdorsal modified type 2 glandular cell outlet; sdrss, subdorsal rounded sensory spot; slne, sublateral nephridiopore; sls, sublateral spine; te, tergal extension; tsp, trichoscalid plate; vldss, ventrolateral droplet-shaped sensory spot; vmdss, ventromedial droplet-shaped sensory spot; vmgco1, ventromedial type 1 glandular cell outlet; vmrss, ventromedial rounded sensory spot." figureDoi="http://doi.org/10.5281/zenodo.10375299" httpUri="https://zenodo.org/record/10375299/files/figure.png" pageId="3" pageNumber="182">Fig. 3A–D</figureCitation>
;
<figureCitation id="BE9B2A1AC676FF8422F1F91D7CA5F926" box="[1169,1179,1746,1765]" captionStart="Fig" captionStartId="5.[994,1024,149,166]" captionTargetBox="[102,967,150,1427]" captionTargetId="figure-287@5.[100,969,148,1429]" captionTargetPageId="5" captionText="Fig. 5. Light micrographs of female holotype MNHN- 655Ma (E), male paratype MNHN-661Ma (C–D), and additional specimens (A-B, F) of Echinoderes angelae sp. nov. showing trunk cuticular details. A: Middorsal to midlateral view on right side of tergal plate of segments 1–2; B: midlateral to ventromedial view on left side of cuticular plates of segments 5–7; C: ventral view of segments 5–6; D: dorsal view of segments 8–9; E: midlateral to ventromedial view on right side of cuticular plates of segments 9–10 (inset shows the intraspecific variability of the nephridiopore); F: midlateral to ventromedial view on right side of cuticular plates of segments 7–8. Abbreviations: lat, lateral accessory tube; lvt, lateroventral tube; ms, muscular scar; ppf, primary pectinate fringe; slne, sublateral nephridiopore; sls, sublateral spine; numbers after abbreviation indicate corresponding segment; glandular cell outlets are marked with continuous circles, and sensory spots with dashed circles or teardrop-like figures." figureDoi="http://doi.org/10.5281/zenodo.10375305" httpUri="https://zenodo.org/record/10375305/files/figure.png" pageId="3" pageNumber="182">5</figureCitation>
A-F; 7B, D, F, H, L). Posterior segment margins straight, with serrated primary pectinate fringe (
<figureCitation id="BE9B2A1AC676FF84255AF8C57BA0F8DE" box="[826,926,1802,1821]" captionStart="Fig" captionStartId="3.[994,1024,149,166]" captionTargetBox="[101,967,149,1199]" captionTargetId="figure-599@3.[100,969,148,1201]" captionTargetPageId="3" captionText="Fig. 3. Line art drawing of Echinoderes angelae sp. nov. A: Ventral female overview; B: dorsal female overview; C: ventral male overview of segments 10–11; D: dorsal male overview of segments 10–11. Abbreviations: dpl, dorsal placid; go, gonopore; lat, lateral accessory tube; lddss, laterodorsal droplet-shaped sensory spot; ldmgco2, laterodorsal modified type 2 glandular cell outlet; ldrss, laterodorsal rounded sensory spot; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; lvgco1, lateroventral type 1 glandular cell outlet; lvmgco2, lateroventral modified type 2 glandular cell outlet; lvt, lateroventral tube; mddss, middorsal droplet-shaped sensory spot; mdgco1, middorsal type 1 glandular cell outlet; mds, middorsal spine; mldss, midlateral droplet-shaped sensory spot; mlmgco2, midlateral modified type 2 glandular cell outlet; ms, muscular scar; mvpl, midventral placid; ppf, primary pectinate fringe; ps, penile spines; S, segment (followed by number of corresponding segment); sddss, subdorsal droplet-shaped sensory spot; sdgco1, subdorsal type 1 glandular cell outlet; sdmgco2, subdorsal modified type 2 glandular cell outlet; sdrss, subdorsal rounded sensory spot; slne, sublateral nephridiopore; sls, sublateral spine; te, tergal extension; tsp, trichoscalid plate; vldss, ventrolateral droplet-shaped sensory spot; vmdss, ventromedial droplet-shaped sensory spot; vmgco1, ventromedial type 1 glandular cell outlet; vmrss, ventromedial rounded sensory spot." figureDoi="http://doi.org/10.5281/zenodo.10375299" httpUri="https://zenodo.org/record/10375299/files/figure.png" pageId="3" pageNumber="182">Fig. 3A–D</figureCitation>
); primary pectinate fringe of segment 1 with larger tips (
<figureCitation id="BE9B2A1AC676FF84255AF8E97B40F8FA" box="[826,894,1830,1849]" captionStart="Fig" captionStartId="7.[995,1025,149,166]" captionTargetBox="[102,968,150,1247]" captionTargetId="figure-378@7.[100,970,148,1249]" captionTargetPageId="7" captionText="Fig. 7. Scanning electron micrographs of additional specimens of Echinoderes angelae sp. nov. showing trunk segments and details of cuticular appendages. A: Subdorsal and laterodorsal sensory spots of segment 1, with the enlarged external ring of micropapillar (inset shows the intraspecific variability of these structures without such kind of external ring); B: ventral view of segments 2–6; C: subdorsal (top) and laterodorsal (bottom) modified type 2 glandular cell outlets of segments 2 and 4 respectively; D: laterodorsal view on left side of tergal plate of segment 2; E: nephridiopore and midlateral dropletshaped sensory spot of segment 9; F: subdorsal to midlateral view on right side of tergal plates of segments 8–9; G: middorsal spine of segment 4; H: subdorsal to midlateral view on left side of tergal plates of segments 3–4; I: laterodorsal tube of segment 10; J: lateral accessory tube of segment 8 (left) and sublateral spine of segment 7 (right); K: subdorsal type 1 glandular cell outlet of segment 9; L: ventral view of segments 10–11; M: subdorsal droplet-shaped sensory spot of segment 9. Abbreviations: go, gonopore; ms, muscular scar; spf, secondary pectinate fringe; modified type 2 glandular cell outlets are marked with continuous circles, and sensory spots with dashed teardrop-like figures." figureDoi="http://doi.org/10.5281/zenodo.10375309" httpUri="https://zenodo.org/record/10375309/files/figure.png" pageId="3" pageNumber="182">Fig. 7B</figureCitation>
); segments 2 and 10 with posterior segment margin extended as a triangle in the ventrolateral to ventromedial region, with tips of primary pectinate fringe conspicuously shorter and tighter than those on other regions on the same segment (
<figureCitation id="BE9B2A1AC676FF84221DF8B57CE4F84E" box="[1149,1242,1914,1933]" captionStart="Fig" captionStartId="3.[994,1024,149,166]" captionTargetBox="[101,967,149,1199]" captionTargetId="figure-599@3.[100,969,148,1201]" captionTargetPageId="3" captionText="Fig. 3. Line art drawing of Echinoderes angelae sp. nov. A: Ventral female overview; B: dorsal female overview; C: ventral male overview of segments 10–11; D: dorsal male overview of segments 10–11. Abbreviations: dpl, dorsal placid; go, gonopore; lat, lateral accessory tube; lddss, laterodorsal droplet-shaped sensory spot; ldmgco2, laterodorsal modified type 2 glandular cell outlet; ldrss, laterodorsal rounded sensory spot; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; lvgco1, lateroventral type 1 glandular cell outlet; lvmgco2, lateroventral modified type 2 glandular cell outlet; lvt, lateroventral tube; mddss, middorsal droplet-shaped sensory spot; mdgco1, middorsal type 1 glandular cell outlet; mds, middorsal spine; mldss, midlateral droplet-shaped sensory spot; mlmgco2, midlateral modified type 2 glandular cell outlet; ms, muscular scar; mvpl, midventral placid; ppf, primary pectinate fringe; ps, penile spines; S, segment (followed by number of corresponding segment); sddss, subdorsal droplet-shaped sensory spot; sdgco1, subdorsal type 1 glandular cell outlet; sdmgco2, subdorsal modified type 2 glandular cell outlet; sdrss, subdorsal rounded sensory spot; slne, sublateral nephridiopore; sls, sublateral spine; te, tergal extension; tsp, trichoscalid plate; vldss, ventrolateral droplet-shaped sensory spot; vmdss, ventromedial droplet-shaped sensory spot; vmgco1, ventromedial type 1 glandular cell outlet; vmrss, ventromedial rounded sensory spot." figureDoi="http://doi.org/10.5281/zenodo.10375299" httpUri="https://zenodo.org/record/10375299/files/figure.png" pageId="3" pageNumber="182">Fig. 3A–D</figureCitation>
;
<figureCitation id="BE9B2A1AC676FF842282F8B57D2AF84E" box="[1250,1300,1914,1933]" captionStart="Fig" captionStartId="7.[995,1025,149,166]" captionTargetBox="[102,968,150,1247]" captionTargetId="figure-378@7.[100,970,148,1249]" captionTargetPageId="7" captionText="Fig. 7. Scanning electron micrographs of additional specimens of Echinoderes angelae sp. nov. showing trunk segments and details of cuticular appendages. A: Subdorsal and laterodorsal sensory spots of segment 1, with the enlarged external ring of micropapillar (inset shows the intraspecific variability of these structures without such kind of external ring); B: ventral view of segments 2–6; C: subdorsal (top) and laterodorsal (bottom) modified type 2 glandular cell outlets of segments 2 and 4 respectively; D: laterodorsal view on left side of tergal plate of segment 2; E: nephridiopore and midlateral dropletshaped sensory spot of segment 9; F: subdorsal to midlateral view on right side of tergal plates of segments 8–9; G: middorsal spine of segment 4; H: subdorsal to midlateral view on left side of tergal plates of segments 3–4; I: laterodorsal tube of segment 10; J: lateral accessory tube of segment 8 (left) and sublateral spine of segment 7 (right); K: subdorsal type 1 glandular cell outlet of segment 9; L: ventral view of segments 10–11; M: subdorsal droplet-shaped sensory spot of segment 9. Abbreviations: go, gonopore; ms, muscular scar; spf, secondary pectinate fringe; modified type 2 glandular cell outlets are marked with continuous circles, and sensory spots with dashed teardrop-like figures." figureDoi="http://doi.org/10.5281/zenodo.10375309" httpUri="https://zenodo.org/record/10375309/files/figure.png" pageId="3" pageNumber="182">7B, L</figureCitation>
); segments 3–9 with tips of primary pectinate fringes noticeable shorter and tighter in the ventromedial region than those on other regions on the same segment (
<figureCitation id="BE9B2A1AC671FF83260CFB82788FFBA3" box="[108,177,1101,1121]" captionStart="Fig" captionStartId="7.[995,1025,149,166]" captionTargetBox="[102,968,150,1247]" captionTargetId="figure-378@7.[100,970,148,1249]" captionTargetPageId="7" captionText="Fig. 7. Scanning electron micrographs of additional specimens of Echinoderes angelae sp. nov. showing trunk segments and details of cuticular appendages. A: Subdorsal and laterodorsal sensory spots of segment 1, with the enlarged external ring of micropapillar (inset shows the intraspecific variability of these structures without such kind of external ring); B: ventral view of segments 2–6; C: subdorsal (top) and laterodorsal (bottom) modified type 2 glandular cell outlets of segments 2 and 4 respectively; D: laterodorsal view on left side of tergal plate of segment 2; E: nephridiopore and midlateral dropletshaped sensory spot of segment 9; F: subdorsal to midlateral view on right side of tergal plates of segments 8–9; G: middorsal spine of segment 4; H: subdorsal to midlateral view on left side of tergal plates of segments 3–4; I: laterodorsal tube of segment 10; J: lateral accessory tube of segment 8 (left) and sublateral spine of segment 7 (right); K: subdorsal type 1 glandular cell outlet of segment 9; L: ventral view of segments 10–11; M: subdorsal droplet-shaped sensory spot of segment 9. Abbreviations: go, gonopore; ms, muscular scar; spf, secondary pectinate fringe; modified type 2 glandular cell outlets are marked with continuous circles, and sensory spots with dashed teardrop-like figures." figureDoi="http://doi.org/10.5281/zenodo.10375309" httpUri="https://zenodo.org/record/10375309/files/figure.png" pageId="4" pageNumber="183">Fig. 7B</figureCitation>
); segment 11 with elongated tips of primary pectinate fringe in middorsal to subdorsal and lateroventral to ventrolateral regions (
<figureCitation id="BE9B2A1AC671FF83260CFB4A78F5FB5B" box="[108,203,1157,1176]" captionStart="Fig" captionStartId="3.[994,1024,149,166]" captionTargetBox="[101,967,149,1199]" captionTargetId="figure-599@3.[100,969,148,1201]" captionTargetPageId="3" captionText="Fig. 3. Line art drawing of Echinoderes angelae sp. nov. A: Ventral female overview; B: dorsal female overview; C: ventral male overview of segments 10–11; D: dorsal male overview of segments 10–11. Abbreviations: dpl, dorsal placid; go, gonopore; lat, lateral accessory tube; lddss, laterodorsal droplet-shaped sensory spot; ldmgco2, laterodorsal modified type 2 glandular cell outlet; ldrss, laterodorsal rounded sensory spot; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; lvgco1, lateroventral type 1 glandular cell outlet; lvmgco2, lateroventral modified type 2 glandular cell outlet; lvt, lateroventral tube; mddss, middorsal droplet-shaped sensory spot; mdgco1, middorsal type 1 glandular cell outlet; mds, middorsal spine; mldss, midlateral droplet-shaped sensory spot; mlmgco2, midlateral modified type 2 glandular cell outlet; ms, muscular scar; mvpl, midventral placid; ppf, primary pectinate fringe; ps, penile spines; S, segment (followed by number of corresponding segment); sddss, subdorsal droplet-shaped sensory spot; sdgco1, subdorsal type 1 glandular cell outlet; sdmgco2, subdorsal modified type 2 glandular cell outlet; sdrss, subdorsal rounded sensory spot; slne, sublateral nephridiopore; sls, sublateral spine; te, tergal extension; tsp, trichoscalid plate; vldss, ventrolateral droplet-shaped sensory spot; vmdss, ventromedial droplet-shaped sensory spot; vmgco1, ventromedial type 1 glandular cell outlet; vmrss, ventromedial rounded sensory spot." figureDoi="http://doi.org/10.5281/zenodo.10375299" httpUri="https://zenodo.org/record/10375299/files/figure.png" pageId="4" pageNumber="183">Fig. 3A–D</figureCitation>
;
<figureCitation id="BE9B2A1AC671FF8326B4FB4A78E1FB5B" box="[212,223,1157,1176]" captionStart="Fig" captionStartId="6.[997,1027,149,166]" captionTargetBox="[102,970,150,1260]" captionTargetId="figure-385@6.[100,972,148,1261]" captionTargetPageId="6" captionText="Fig. 6. Scanning electron micrographs of additional specimens of Echinoderes angelae sp. nov. showing trunk overview and details of introvert, neck, lateral terminal spines and sexually dimorphic features. A: Lateral trunk overview; B: introvert, sector 3; C: introvert, sector 4; D: ventral neck view; E: lateral terminal spine; F: male penile spines; G: female lateral terminal accessory spine. Abbreviations: ba, bacteria; ftz, fringed transverse zone; ltas, lateral terminal accessory spine; lts, lateral terminal spine; mvpl, midventral placid; ne, neck; ps, penile spines; psp, primary spinoscalid; sp, spinoscalid (followed by number of corresponding ring); te, tergal extension; ts, trichoscalid; tsp, trichoscalid plate; lambda marks (̂) indicate attachment point of spinoscalids." figureDoi="http://doi.org/10.5281/zenodo.10375307" httpUri="https://zenodo.org/record/10375307/files/figure.png" pageId="4" pageNumber="183">6</figureCitation>
F-G). Secondary pectinate fringes finely serrated (
<figureCitation id="BE9B2A1AC671FF8324D2FB4A7AC6FB5B" box="[690,760,1157,1176]" captionStart="Fig" captionStartId="7.[995,1025,149,166]" captionTargetBox="[102,968,150,1247]" captionTargetId="figure-378@7.[100,970,148,1249]" captionTargetPageId="7" captionText="Fig. 7. Scanning electron micrographs of additional specimens of Echinoderes angelae sp. nov. showing trunk segments and details of cuticular appendages. A: Subdorsal and laterodorsal sensory spots of segment 1, with the enlarged external ring of micropapillar (inset shows the intraspecific variability of these structures without such kind of external ring); B: ventral view of segments 2–6; C: subdorsal (top) and laterodorsal (bottom) modified type 2 glandular cell outlets of segments 2 and 4 respectively; D: laterodorsal view on left side of tergal plate of segment 2; E: nephridiopore and midlateral dropletshaped sensory spot of segment 9; F: subdorsal to midlateral view on right side of tergal plates of segments 8–9; G: middorsal spine of segment 4; H: subdorsal to midlateral view on left side of tergal plates of segments 3–4; I: laterodorsal tube of segment 10; J: lateral accessory tube of segment 8 (left) and sublateral spine of segment 7 (right); K: subdorsal type 1 glandular cell outlet of segment 9; L: ventral view of segments 10–11; M: subdorsal droplet-shaped sensory spot of segment 9. Abbreviations: go, gonopore; ms, muscular scar; spf, secondary pectinate fringe; modified type 2 glandular cell outlets are marked with continuous circles, and sensory spots with dashed teardrop-like figures." figureDoi="http://doi.org/10.5281/zenodo.10375309" httpUri="https://zenodo.org/record/10375309/files/figure.png" pageId="4" pageNumber="183">Fig. 7K</figureCitation>
).
</paragraph>
<caption id="72DF6617C671FF832583FF5A7DAEFDFC" ID-DOI="http://doi.org/10.5281/zenodo.10375301" ID-Zenodo-Dep="10375301" httpUri="https://zenodo.org/record/10375301/files/figure.png" pageId="4" pageNumber="183" startId="4.[995,1025,149,166]" targetBox="[102,968,150,977]" targetPageId="4" targetType="figure">
<paragraph id="261F369FC671FF832583FF5A7DAEFDFC" blockId="4.[995,1488,149,575]" pageId="4" pageNumber="183">
<emphasis id="14D4EA8DC671FF832583FF5A7C1CFF65" bold="true" box="[995,1058,149,166]" pageId="4" pageNumber="183">Fig. 4.</emphasis>
Light micrographs of female holotype MNHN- 655Ma (A, C-D), male paratype MNHN-661Ma (B, E), female paratypes MNHN-656Ma (H) and MNHN-657Ma (F), and additional specimens (G, I) of
<taxonomicName id="E1A04D1CC671FF83236FFF2D7D8FFF30" authority="Cepeda &amp; Gayet &amp; Spedicato &amp; Michaud &amp; Zeppilli, 2022" authorityName="Cepeda &amp; Gayet &amp; Spedicato &amp; Michaud &amp; Zeppilli" authorityYear="2022" box="[1295,1457,226,243]" class="Cyclorhagida" family="Echinoderidae" genus="Echinoderes" kingdom="Animalia" order="Echinorhagata" pageId="4" pageNumber="183" phylum="Kinorhyncha" rank="species" species="angelae" status="sp. nov.">
<emphasis id="14D4EA8DC671FF83236FFF2D7D8FFF30" bold="true" box="[1295,1457,226,243]" italics="true" pageId="4" pageNumber="183">Echinoderes angelae</emphasis>
</taxonomicName>
<taxonomicNameLabel id="0FE757F6C671FF8323D7FF2D7C37FECF" pageId="4" pageNumber="183" rank="species">sp. nov.</taxonomicNameLabel>
showing trunk overview, details of mouth cone and neck, epibionts and ingested diatoms. A: Ventral trunk overview; B: mouth cone, showing the outer oral styles; C: dorsal neck view; D: ventral neck view; E: middorsal to laterodorsal view on right side of tergal plate of segments 10–11; F: lateral view of segment 11; G: ingested diatom; H: unknown type of epibionts attached to the posterior edge of segment 11; I: filamentous bacteria attached to the posterior edge of segment 11. Abbreviations: ba, bacteria; bs, basal sheath; dpl, dorsal placid; ep, end-piece; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; mvpl, midventral placid; ps, penile spines; te, tergal extension; tsp, trichoscalid plate.
</paragraph>
</caption>
<paragraph id="261F369FC671FF8326E4FB6E7AE9FAB4" blockId="4.[100,771,1073,1957]" pageId="4" pageNumber="183">
<materialsCitation id="96C83CC2C671FF8326E4FB6E7AEDFAB4" collectionCode="SEM" pageId="4" pageNumber="183" specimenCount="7">
Segment 1 with
<typeStatus id="F91B883DC671FF832742FB6E7975FB77" box="[290,331,1185,1204]" pageId="4" pageNumber="183">type</typeStatus>
1 glandular cell outlets in middorsal and lateroventral positions (
<figureCitation id="BE9B2A1AC671FF83274BFB727996FB13" box="[299,424,1213,1232]" captionStart="Fig" captionStartId="3.[994,1024,149,166]" captionTargetBox="[101,967,149,1199]" captionTargetId="figure-599@3.[100,969,148,1201]" captionTargetPageId="3" captionText="Fig. 3. Line art drawing of Echinoderes angelae sp. nov. A: Ventral female overview; B: dorsal female overview; C: ventral male overview of segments 10–11; D: dorsal male overview of segments 10–11. Abbreviations: dpl, dorsal placid; go, gonopore; lat, lateral accessory tube; lddss, laterodorsal droplet-shaped sensory spot; ldmgco2, laterodorsal modified type 2 glandular cell outlet; ldrss, laterodorsal rounded sensory spot; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; lvgco1, lateroventral type 1 glandular cell outlet; lvmgco2, lateroventral modified type 2 glandular cell outlet; lvt, lateroventral tube; mddss, middorsal droplet-shaped sensory spot; mdgco1, middorsal type 1 glandular cell outlet; mds, middorsal spine; mldss, midlateral droplet-shaped sensory spot; mlmgco2, midlateral modified type 2 glandular cell outlet; ms, muscular scar; mvpl, midventral placid; ppf, primary pectinate fringe; ps, penile spines; S, segment (followed by number of corresponding segment); sddss, subdorsal droplet-shaped sensory spot; sdgco1, subdorsal type 1 glandular cell outlet; sdmgco2, subdorsal modified type 2 glandular cell outlet; sdrss, subdorsal rounded sensory spot; slne, sublateral nephridiopore; sls, sublateral spine; te, tergal extension; tsp, trichoscalid plate; vldss, ventrolateral droplet-shaped sensory spot; vmdss, ventromedial droplet-shaped sensory spot; vmgco1, ventromedial type 1 glandular cell outlet; vmrss, ventromedial rounded sensory spot." figureDoi="http://doi.org/10.5281/zenodo.10375299" httpUri="https://zenodo.org/record/10375299/files/figure.png" pageId="4" pageNumber="183">Fig. 3A and B</figureCitation>
;
<figureCitation id="BE9B2A1AC671FF8327D1FB7279F1FB13" box="[433,463,1213,1232]" captionStart="Fig" captionStartId="5.[994,1024,149,166]" captionTargetBox="[102,967,150,1427]" captionTargetId="figure-287@5.[100,969,148,1429]" captionTargetPageId="5" captionText="Fig. 5. Light micrographs of female holotype MNHN- 655Ma (E), male paratype MNHN-661Ma (C–D), and additional specimens (A-B, F) of Echinoderes angelae sp. nov. showing trunk cuticular details. A: Middorsal to midlateral view on right side of tergal plate of segments 1–2; B: midlateral to ventromedial view on left side of cuticular plates of segments 5–7; C: ventral view of segments 5–6; D: dorsal view of segments 8–9; E: midlateral to ventromedial view on right side of cuticular plates of segments 9–10 (inset shows the intraspecific variability of the nephridiopore); F: midlateral to ventromedial view on right side of cuticular plates of segments 7–8. Abbreviations: lat, lateral accessory tube; lvt, lateroventral tube; ms, muscular scar; ppf, primary pectinate fringe; slne, sublateral nephridiopore; sls, sublateral spine; numbers after abbreviation indicate corresponding segment; glandular cell outlets are marked with continuous circles, and sensory spots with dashed circles or teardrop-like figures." figureDoi="http://doi.org/10.5281/zenodo.10375305" httpUri="https://zenodo.org/record/10375305/files/figure.png" pageId="4" pageNumber="183">5A</figureCitation>
). Large, rounded sensory spots in subdorsal, laterodorsal and ventromedial positions, the former two located near the anterior segment margin, the latter near the posterior segment edge; these sensory spots have a transversal row of conspicuously elongated hairs at the posterior edge of the papillae area (except in
<specimenCount id="30A6FD16C671FF832604FA86793EFA98" box="[100,256,1352,1372]" count="7" pageId="4" pageNumber="183" type="generic">seven specimens</specimenCount>
mounted for
<collectionCode id="40B1AE5AC671FF8327E4FA877990FA98" box="[388,430,1352,1371]" pageId="4" pageNumber="183">SEM</collectionCode>
that lacked these hairs, see subsection
<emphasis id="14D4EA8DC671FF8326F0FAAB79DBFAB4" bold="true" box="[144,485,1380,1399]" italics="true" pageId="4" pageNumber="183">3.1.5 Notes on intraspecific variability</emphasis>
) (
<figureCitation id="BE9B2A1AC671FF83279BFAAB7A43FABB" box="[507,637,1380,1400]" captionStart="Fig" captionStartId="3.[994,1024,149,166]" captionTargetBox="[101,967,149,1199]" captionTargetId="figure-599@3.[100,969,148,1201]" captionTargetPageId="3" captionText="Fig. 3. Line art drawing of Echinoderes angelae sp. nov. A: Ventral female overview; B: dorsal female overview; C: ventral male overview of segments 10–11; D: dorsal male overview of segments 10–11. Abbreviations: dpl, dorsal placid; go, gonopore; lat, lateral accessory tube; lddss, laterodorsal droplet-shaped sensory spot; ldmgco2, laterodorsal modified type 2 glandular cell outlet; ldrss, laterodorsal rounded sensory spot; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; lvgco1, lateroventral type 1 glandular cell outlet; lvmgco2, lateroventral modified type 2 glandular cell outlet; lvt, lateroventral tube; mddss, middorsal droplet-shaped sensory spot; mdgco1, middorsal type 1 glandular cell outlet; mds, middorsal spine; mldss, midlateral droplet-shaped sensory spot; mlmgco2, midlateral modified type 2 glandular cell outlet; ms, muscular scar; mvpl, midventral placid; ppf, primary pectinate fringe; ps, penile spines; S, segment (followed by number of corresponding segment); sddss, subdorsal droplet-shaped sensory spot; sdgco1, subdorsal type 1 glandular cell outlet; sdmgco2, subdorsal modified type 2 glandular cell outlet; sdrss, subdorsal rounded sensory spot; slne, sublateral nephridiopore; sls, sublateral spine; te, tergal extension; tsp, trichoscalid plate; vldss, ventrolateral droplet-shaped sensory spot; vmdss, ventromedial droplet-shaped sensory spot; vmgco1, ventromedial type 1 glandular cell outlet; vmrss, ventromedial rounded sensory spot." figureDoi="http://doi.org/10.5281/zenodo.10375299" httpUri="https://zenodo.org/record/10375299/files/figure.png" pageId="4" pageNumber="183">Fig. 3A and B</figureCitation>
;
<figureCitation id="BE9B2A1AC671FF8324E8FAAB7A9AFAB4" box="[648,676,1380,1399]" captionStart="Fig" captionStartId="5.[994,1024,149,166]" captionTargetBox="[102,967,150,1427]" captionTargetId="figure-287@5.[100,969,148,1429]" captionTargetPageId="5" captionText="Fig. 5. Light micrographs of female holotype MNHN- 655Ma (E), male paratype MNHN-661Ma (C–D), and additional specimens (A-B, F) of Echinoderes angelae sp. nov. showing trunk cuticular details. A: Middorsal to midlateral view on right side of tergal plate of segments 1–2; B: midlateral to ventromedial view on left side of cuticular plates of segments 5–7; C: ventral view of segments 5–6; D: dorsal view of segments 8–9; E: midlateral to ventromedial view on right side of cuticular plates of segments 9–10 (inset shows the intraspecific variability of the nephridiopore); F: midlateral to ventromedial view on right side of cuticular plates of segments 7–8. Abbreviations: lat, lateral accessory tube; lvt, lateroventral tube; ms, muscular scar; ppf, primary pectinate fringe; slne, sublateral nephridiopore; sls, sublateral spine; numbers after abbreviation indicate corresponding segment; glandular cell outlets are marked with continuous circles, and sensory spots with dashed circles or teardrop-like figures." figureDoi="http://doi.org/10.5281/zenodo.10375305" httpUri="https://zenodo.org/record/10375305/files/figure.png" pageId="4" pageNumber="183">5A</figureCitation>
;
<figureCitation id="BE9B2A1AC671FF8324CEFAAB7AF3FAB4" box="[686,717,1380,1399]" captionStart="Fig" captionStartId="7.[995,1025,149,166]" captionTargetBox="[102,968,150,1247]" captionTargetId="figure-378@7.[100,970,148,1249]" captionTargetPageId="7" captionText="Fig. 7. Scanning electron micrographs of additional specimens of Echinoderes angelae sp. nov. showing trunk segments and details of cuticular appendages. A: Subdorsal and laterodorsal sensory spots of segment 1, with the enlarged external ring of micropapillar (inset shows the intraspecific variability of these structures without such kind of external ring); B: ventral view of segments 2–6; C: subdorsal (top) and laterodorsal (bottom) modified type 2 glandular cell outlets of segments 2 and 4 respectively; D: laterodorsal view on left side of tergal plate of segment 2; E: nephridiopore and midlateral dropletshaped sensory spot of segment 9; F: subdorsal to midlateral view on right side of tergal plates of segments 8–9; G: middorsal spine of segment 4; H: subdorsal to midlateral view on left side of tergal plates of segments 3–4; I: laterodorsal tube of segment 10; J: lateral accessory tube of segment 8 (left) and sublateral spine of segment 7 (right); K: subdorsal type 1 glandular cell outlet of segment 9; L: ventral view of segments 10–11; M: subdorsal droplet-shaped sensory spot of segment 9. Abbreviations: go, gonopore; ms, muscular scar; spf, secondary pectinate fringe; modified type 2 glandular cell outlets are marked with continuous circles, and sensory spots with dashed teardrop-like figures." figureDoi="http://doi.org/10.5281/zenodo.10375309" httpUri="https://zenodo.org/record/10375309/files/figure.png" pageId="4" pageNumber="183">7A</figureCitation>
)
</materialsCitation>
.
</paragraph>
<paragraph id="261F369FC671FF8326E4FA4F7880F9B1" blockId="4.[100,771,1073,1957]" pageId="4" pageNumber="183">
Segment 2 with one pair of droplet-shaped sensory spots in middorsal and ventromedial positions, and two pairs in laterodorsal position (
<figureCitation id="BE9B2A1AC671FF8326A2FA777970FA08" box="[194,334,1464,1483]" captionStart="Fig" captionStartId="3.[994,1024,149,166]" captionTargetBox="[101,967,149,1199]" captionTargetId="figure-599@3.[100,969,148,1201]" captionTargetPageId="3" captionText="Fig. 3. Line art drawing of Echinoderes angelae sp. nov. A: Ventral female overview; B: dorsal female overview; C: ventral male overview of segments 10–11; D: dorsal male overview of segments 10–11. Abbreviations: dpl, dorsal placid; go, gonopore; lat, lateral accessory tube; lddss, laterodorsal droplet-shaped sensory spot; ldmgco2, laterodorsal modified type 2 glandular cell outlet; ldrss, laterodorsal rounded sensory spot; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; lvgco1, lateroventral type 1 glandular cell outlet; lvmgco2, lateroventral modified type 2 glandular cell outlet; lvt, lateroventral tube; mddss, middorsal droplet-shaped sensory spot; mdgco1, middorsal type 1 glandular cell outlet; mds, middorsal spine; mldss, midlateral droplet-shaped sensory spot; mlmgco2, midlateral modified type 2 glandular cell outlet; ms, muscular scar; mvpl, midventral placid; ppf, primary pectinate fringe; ps, penile spines; S, segment (followed by number of corresponding segment); sddss, subdorsal droplet-shaped sensory spot; sdgco1, subdorsal type 1 glandular cell outlet; sdmgco2, subdorsal modified type 2 glandular cell outlet; sdrss, subdorsal rounded sensory spot; slne, sublateral nephridiopore; sls, sublateral spine; te, tergal extension; tsp, trichoscalid plate; vldss, ventrolateral droplet-shaped sensory spot; vmdss, ventromedial droplet-shaped sensory spot; vmgco1, ventromedial type 1 glandular cell outlet; vmrss, ventromedial rounded sensory spot." figureDoi="http://doi.org/10.5281/zenodo.10375299" httpUri="https://zenodo.org/record/10375299/files/figure.png" pageId="4" pageNumber="183">Fig. 3A and B</figureCitation>
;
<figureCitation id="BE9B2A1AC671FF83273CFA777944FA08" box="[348,378,1464,1483]" captionStart="Fig" captionStartId="5.[994,1024,149,166]" captionTargetBox="[102,967,150,1427]" captionTargetId="figure-287@5.[100,969,148,1429]" captionTargetPageId="5" captionText="Fig. 5. Light micrographs of female holotype MNHN- 655Ma (E), male paratype MNHN-661Ma (C–D), and additional specimens (A-B, F) of Echinoderes angelae sp. nov. showing trunk cuticular details. A: Middorsal to midlateral view on right side of tergal plate of segments 1–2; B: midlateral to ventromedial view on left side of cuticular plates of segments 5–7; C: ventral view of segments 5–6; D: dorsal view of segments 8–9; E: midlateral to ventromedial view on right side of cuticular plates of segments 9–10 (inset shows the intraspecific variability of the nephridiopore); F: midlateral to ventromedial view on right side of cuticular plates of segments 7–8. Abbreviations: lat, lateral accessory tube; lvt, lateroventral tube; ms, muscular scar; ppf, primary pectinate fringe; slne, sublateral nephridiopore; sls, sublateral spine; numbers after abbreviation indicate corresponding segment; glandular cell outlets are marked with continuous circles, and sensory spots with dashed circles or teardrop-like figures." figureDoi="http://doi.org/10.5281/zenodo.10375305" httpUri="https://zenodo.org/record/10375305/files/figure.png" pageId="4" pageNumber="183">5A</figureCitation>
); on this and following segments, these sensory spots usually have one or two central pores with emerging cilia (
<figureCitation id="BE9B2A1AC671FF83260CFA207886F9C0" box="[108,184,1519,1539]" captionStart="Fig" captionStartId="7.[995,1025,149,166]" captionTargetBox="[102,968,150,1247]" captionTargetId="figure-378@7.[100,970,148,1249]" captionTargetPageId="7" captionText="Fig. 7. Scanning electron micrographs of additional specimens of Echinoderes angelae sp. nov. showing trunk segments and details of cuticular appendages. A: Subdorsal and laterodorsal sensory spots of segment 1, with the enlarged external ring of micropapillar (inset shows the intraspecific variability of these structures without such kind of external ring); B: ventral view of segments 2–6; C: subdorsal (top) and laterodorsal (bottom) modified type 2 glandular cell outlets of segments 2 and 4 respectively; D: laterodorsal view on left side of tergal plate of segment 2; E: nephridiopore and midlateral dropletshaped sensory spot of segment 9; F: subdorsal to midlateral view on right side of tergal plates of segments 8–9; G: middorsal spine of segment 4; H: subdorsal to midlateral view on left side of tergal plates of segments 3–4; I: laterodorsal tube of segment 10; J: lateral accessory tube of segment 8 (left) and sublateral spine of segment 7 (right); K: subdorsal type 1 glandular cell outlet of segment 9; L: ventral view of segments 10–11; M: subdorsal droplet-shaped sensory spot of segment 9. Abbreviations: go, gonopore; ms, muscular scar; spf, secondary pectinate fringe; modified type 2 glandular cell outlets are marked with continuous circles, and sensory spots with dashed teardrop-like figures." figureDoi="http://doi.org/10.5281/zenodo.10375309" httpUri="https://zenodo.org/record/10375309/files/figure.png" pageId="4" pageNumber="183">Fig. 7D</figureCitation>
). Modified
<typeStatus id="F91B883DC671FF83274AFA3F796DF9C0" box="[298,339,1520,1539]" pageId="4" pageNumber="183">type</typeStatus>
2 glandular cell outlets in subdorsal, laterodorsal and midlateral positions (
<figureCitation id="BE9B2A1AC671FF8327FDF9C479DFF9DC" box="[413,481,1547,1567]" captionStart="Fig" captionStartId="3.[994,1024,149,166]" captionTargetBox="[101,967,149,1199]" captionTargetId="figure-599@3.[100,969,148,1201]" captionTargetPageId="3" captionText="Fig. 3. Line art drawing of Echinoderes angelae sp. nov. A: Ventral female overview; B: dorsal female overview; C: ventral male overview of segments 10–11; D: dorsal male overview of segments 10–11. Abbreviations: dpl, dorsal placid; go, gonopore; lat, lateral accessory tube; lddss, laterodorsal droplet-shaped sensory spot; ldmgco2, laterodorsal modified type 2 glandular cell outlet; ldrss, laterodorsal rounded sensory spot; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; lvgco1, lateroventral type 1 glandular cell outlet; lvmgco2, lateroventral modified type 2 glandular cell outlet; lvt, lateroventral tube; mddss, middorsal droplet-shaped sensory spot; mdgco1, middorsal type 1 glandular cell outlet; mds, middorsal spine; mldss, midlateral droplet-shaped sensory spot; mlmgco2, midlateral modified type 2 glandular cell outlet; ms, muscular scar; mvpl, midventral placid; ppf, primary pectinate fringe; ps, penile spines; S, segment (followed by number of corresponding segment); sddss, subdorsal droplet-shaped sensory spot; sdgco1, subdorsal type 1 glandular cell outlet; sdmgco2, subdorsal modified type 2 glandular cell outlet; sdrss, subdorsal rounded sensory spot; slne, sublateral nephridiopore; sls, sublateral spine; te, tergal extension; tsp, trichoscalid plate; vldss, ventrolateral droplet-shaped sensory spot; vmdss, ventromedial droplet-shaped sensory spot; vmgco1, ventromedial type 1 glandular cell outlet; vmrss, ventromedial rounded sensory spot." figureDoi="http://doi.org/10.5281/zenodo.10375299" httpUri="https://zenodo.org/record/10375299/files/figure.png" pageId="4" pageNumber="183">Fig. 3B</figureCitation>
;
<figureCitation id="BE9B2A1AC671FF83278AF9C47A38F9DD" box="[490,518,1547,1566]" captionStart="Fig" captionStartId="5.[994,1024,149,166]" captionTargetBox="[102,967,150,1427]" captionTargetId="figure-287@5.[100,969,148,1429]" captionTargetPageId="5" captionText="Fig. 5. Light micrographs of female holotype MNHN- 655Ma (E), male paratype MNHN-661Ma (C–D), and additional specimens (A-B, F) of Echinoderes angelae sp. nov. showing trunk cuticular details. A: Middorsal to midlateral view on right side of tergal plate of segments 1–2; B: midlateral to ventromedial view on left side of cuticular plates of segments 5–7; C: ventral view of segments 5–6; D: dorsal view of segments 8–9; E: midlateral to ventromedial view on right side of cuticular plates of segments 9–10 (inset shows the intraspecific variability of the nephridiopore); F: midlateral to ventromedial view on right side of cuticular plates of segments 7–8. Abbreviations: lat, lateral accessory tube; lvt, lateroventral tube; ms, muscular scar; ppf, primary pectinate fringe; slne, sublateral nephridiopore; sls, sublateral spine; numbers after abbreviation indicate corresponding segment; glandular cell outlets are marked with continuous circles, and sensory spots with dashed circles or teardrop-like figures." figureDoi="http://doi.org/10.5281/zenodo.10375305" httpUri="https://zenodo.org/record/10375305/files/figure.png" pageId="4" pageNumber="183">5A</figureCitation>
;
<figureCitation id="BE9B2A1AC671FF83246FF9C37A13F9DC" box="[527,557,1548,1567]" captionStart="Fig" captionStartId="7.[995,1025,149,166]" captionTargetBox="[102,968,150,1247]" captionTargetId="figure-378@7.[100,970,148,1249]" captionTargetPageId="7" captionText="Fig. 7. Scanning electron micrographs of additional specimens of Echinoderes angelae sp. nov. showing trunk segments and details of cuticular appendages. A: Subdorsal and laterodorsal sensory spots of segment 1, with the enlarged external ring of micropapillar (inset shows the intraspecific variability of these structures without such kind of external ring); B: ventral view of segments 2–6; C: subdorsal (top) and laterodorsal (bottom) modified type 2 glandular cell outlets of segments 2 and 4 respectively; D: laterodorsal view on left side of tergal plate of segment 2; E: nephridiopore and midlateral dropletshaped sensory spot of segment 9; F: subdorsal to midlateral view on right side of tergal plates of segments 8–9; G: middorsal spine of segment 4; H: subdorsal to midlateral view on left side of tergal plates of segments 3–4; I: laterodorsal tube of segment 10; J: lateral accessory tube of segment 8 (left) and sublateral spine of segment 7 (right); K: subdorsal type 1 glandular cell outlet of segment 9; L: ventral view of segments 10–11; M: subdorsal droplet-shaped sensory spot of segment 9. Abbreviations: go, gonopore; ms, muscular scar; spf, secondary pectinate fringe; modified type 2 glandular cell outlets are marked with continuous circles, and sensory spots with dashed teardrop-like figures." figureDoi="http://doi.org/10.5281/zenodo.10375309" httpUri="https://zenodo.org/record/10375309/files/figure.png" pageId="4" pageNumber="183">7D</figureCitation>
); on this and following segments, modified
<typeStatus id="F91B883DC671FF83274CF9E7796BF9F8" box="[300,341,1576,1595]" pageId="4" pageNumber="183">type</typeStatus>
2 glandular cell outlets are minute, oval, glandular openings surrounded by a tuft of short cuticular extensions (
<figureCitation id="BE9B2A1AC671FF83260CF990788DF9B1" box="[108,179,1631,1650]" captionStart="Fig" captionStartId="7.[995,1025,149,166]" captionTargetBox="[102,968,150,1247]" captionTargetId="figure-378@7.[100,970,148,1249]" captionTargetPageId="7" captionText="Fig. 7. Scanning electron micrographs of additional specimens of Echinoderes angelae sp. nov. showing trunk segments and details of cuticular appendages. A: Subdorsal and laterodorsal sensory spots of segment 1, with the enlarged external ring of micropapillar (inset shows the intraspecific variability of these structures without such kind of external ring); B: ventral view of segments 2–6; C: subdorsal (top) and laterodorsal (bottom) modified type 2 glandular cell outlets of segments 2 and 4 respectively; D: laterodorsal view on left side of tergal plate of segment 2; E: nephridiopore and midlateral dropletshaped sensory spot of segment 9; F: subdorsal to midlateral view on right side of tergal plates of segments 8–9; G: middorsal spine of segment 4; H: subdorsal to midlateral view on left side of tergal plates of segments 3–4; I: laterodorsal tube of segment 10; J: lateral accessory tube of segment 8 (left) and sublateral spine of segment 7 (right); K: subdorsal type 1 glandular cell outlet of segment 9; L: ventral view of segments 10–11; M: subdorsal droplet-shaped sensory spot of segment 9. Abbreviations: go, gonopore; ms, muscular scar; spf, secondary pectinate fringe; modified type 2 glandular cell outlets are marked with continuous circles, and sensory spots with dashed teardrop-like figures." figureDoi="http://doi.org/10.5281/zenodo.10375309" httpUri="https://zenodo.org/record/10375309/files/figure.png" pageId="4" pageNumber="183">Fig. 7C</figureCitation>
).
</paragraph>
<paragraph id="261F369FC671FF8326E4F9B47A0EF905" blockId="4.[100,771,1073,1957]" pageId="4" pageNumber="183">
Segment 3 with
<typeStatus id="F91B883DC671FF83274EF9B47969F94D" box="[302,343,1659,1678]" pageId="4" pageNumber="183">type</typeStatus>
1 glandular cell outlets in middorsal and ventromedial positions (
<figureCitation id="BE9B2A1AC671FF832728F95879F7F969" box="[328,457,1687,1706]" captionStart="Fig" captionStartId="3.[994,1024,149,166]" captionTargetBox="[101,967,149,1199]" captionTargetId="figure-599@3.[100,969,148,1201]" captionTargetPageId="3" captionText="Fig. 3. Line art drawing of Echinoderes angelae sp. nov. A: Ventral female overview; B: dorsal female overview; C: ventral male overview of segments 10–11; D: dorsal male overview of segments 10–11. Abbreviations: dpl, dorsal placid; go, gonopore; lat, lateral accessory tube; lddss, laterodorsal droplet-shaped sensory spot; ldmgco2, laterodorsal modified type 2 glandular cell outlet; ldrss, laterodorsal rounded sensory spot; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; lvgco1, lateroventral type 1 glandular cell outlet; lvmgco2, lateroventral modified type 2 glandular cell outlet; lvt, lateroventral tube; mddss, middorsal droplet-shaped sensory spot; mdgco1, middorsal type 1 glandular cell outlet; mds, middorsal spine; mldss, midlateral droplet-shaped sensory spot; mlmgco2, midlateral modified type 2 glandular cell outlet; ms, muscular scar; mvpl, midventral placid; ppf, primary pectinate fringe; ps, penile spines; S, segment (followed by number of corresponding segment); sddss, subdorsal droplet-shaped sensory spot; sdgco1, subdorsal type 1 glandular cell outlet; sdmgco2, subdorsal modified type 2 glandular cell outlet; sdrss, subdorsal rounded sensory spot; slne, sublateral nephridiopore; sls, sublateral spine; te, tergal extension; tsp, trichoscalid plate; vldss, ventrolateral droplet-shaped sensory spot; vmdss, ventromedial droplet-shaped sensory spot; vmgco1, ventromedial type 1 glandular cell outlet; vmrss, ventromedial rounded sensory spot." figureDoi="http://doi.org/10.5281/zenodo.10375299" httpUri="https://zenodo.org/record/10375299/files/figure.png" pageId="4" pageNumber="183">Fig. 3A and B</figureCitation>
). Droplet-shaped sensory spots in subdorsal and midlateral positions (
<figureCitation id="BE9B2A1AC671FF8327D7F97C79C2F905" box="[439,508,1715,1734]" captionStart="Fig" captionStartId="3.[994,1024,149,166]" captionTargetBox="[101,967,149,1199]" captionTargetId="figure-599@3.[100,969,148,1201]" captionTargetPageId="3" captionText="Fig. 3. Line art drawing of Echinoderes angelae sp. nov. A: Ventral female overview; B: dorsal female overview; C: ventral male overview of segments 10–11; D: dorsal male overview of segments 10–11. Abbreviations: dpl, dorsal placid; go, gonopore; lat, lateral accessory tube; lddss, laterodorsal droplet-shaped sensory spot; ldmgco2, laterodorsal modified type 2 glandular cell outlet; ldrss, laterodorsal rounded sensory spot; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; lvgco1, lateroventral type 1 glandular cell outlet; lvmgco2, lateroventral modified type 2 glandular cell outlet; lvt, lateroventral tube; mddss, middorsal droplet-shaped sensory spot; mdgco1, middorsal type 1 glandular cell outlet; mds, middorsal spine; mldss, midlateral droplet-shaped sensory spot; mlmgco2, midlateral modified type 2 glandular cell outlet; ms, muscular scar; mvpl, midventral placid; ppf, primary pectinate fringe; ps, penile spines; S, segment (followed by number of corresponding segment); sddss, subdorsal droplet-shaped sensory spot; sdgco1, subdorsal type 1 glandular cell outlet; sdmgco2, subdorsal modified type 2 glandular cell outlet; sdrss, subdorsal rounded sensory spot; slne, sublateral nephridiopore; sls, sublateral spine; te, tergal extension; tsp, trichoscalid plate; vldss, ventrolateral droplet-shaped sensory spot; vmdss, ventromedial droplet-shaped sensory spot; vmgco1, ventromedial type 1 glandular cell outlet; vmrss, ventromedial rounded sensory spot." figureDoi="http://doi.org/10.5281/zenodo.10375299" httpUri="https://zenodo.org/record/10375299/files/figure.png" pageId="4" pageNumber="183">Fig. 3B</figureCitation>
;
<figureCitation id="BE9B2A1AC671FF832466F97C7A1BF905" box="[518,549,1715,1734]" captionStart="Fig" captionStartId="7.[995,1025,149,166]" captionTargetBox="[102,968,150,1247]" captionTargetId="figure-378@7.[100,970,148,1249]" captionTargetPageId="7" captionText="Fig. 7. Scanning electron micrographs of additional specimens of Echinoderes angelae sp. nov. showing trunk segments and details of cuticular appendages. A: Subdorsal and laterodorsal sensory spots of segment 1, with the enlarged external ring of micropapillar (inset shows the intraspecific variability of these structures without such kind of external ring); B: ventral view of segments 2–6; C: subdorsal (top) and laterodorsal (bottom) modified type 2 glandular cell outlets of segments 2 and 4 respectively; D: laterodorsal view on left side of tergal plate of segment 2; E: nephridiopore and midlateral dropletshaped sensory spot of segment 9; F: subdorsal to midlateral view on right side of tergal plates of segments 8–9; G: middorsal spine of segment 4; H: subdorsal to midlateral view on left side of tergal plates of segments 3–4; I: laterodorsal tube of segment 10; J: lateral accessory tube of segment 8 (left) and sublateral spine of segment 7 (right); K: subdorsal type 1 glandular cell outlet of segment 9; L: ventral view of segments 10–11; M: subdorsal droplet-shaped sensory spot of segment 9. Abbreviations: go, gonopore; ms, muscular scar; spf, secondary pectinate fringe; modified type 2 glandular cell outlets are marked with continuous circles, and sensory spots with dashed teardrop-like figures." figureDoi="http://doi.org/10.5281/zenodo.10375309" httpUri="https://zenodo.org/record/10375309/files/figure.png" pageId="4" pageNumber="183">7H</figureCitation>
).
</paragraph>
<paragraph id="261F369FC671FF8326E4F90078DDF892" blockId="4.[100,771,1073,1957]" pageId="4" pageNumber="183">
Segment 4 with short (ca. 4–8 μm length), poorly sclerotized, weakly articulated, acicular spine in middorsal position (
<figureCitation id="BE9B2A1AC671FF83245EF9247ABAF93D" box="[574,644,1771,1790]" captionStart="Fig" captionStartId="3.[994,1024,149,166]" captionTargetBox="[101,967,149,1199]" captionTargetId="figure-599@3.[100,969,148,1201]" captionTargetPageId="3" captionText="Fig. 3. Line art drawing of Echinoderes angelae sp. nov. A: Ventral female overview; B: dorsal female overview; C: ventral male overview of segments 10–11; D: dorsal male overview of segments 10–11. Abbreviations: dpl, dorsal placid; go, gonopore; lat, lateral accessory tube; lddss, laterodorsal droplet-shaped sensory spot; ldmgco2, laterodorsal modified type 2 glandular cell outlet; ldrss, laterodorsal rounded sensory spot; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; lvgco1, lateroventral type 1 glandular cell outlet; lvmgco2, lateroventral modified type 2 glandular cell outlet; lvt, lateroventral tube; mddss, middorsal droplet-shaped sensory spot; mdgco1, middorsal type 1 glandular cell outlet; mds, middorsal spine; mldss, midlateral droplet-shaped sensory spot; mlmgco2, midlateral modified type 2 glandular cell outlet; ms, muscular scar; mvpl, midventral placid; ppf, primary pectinate fringe; ps, penile spines; S, segment (followed by number of corresponding segment); sddss, subdorsal droplet-shaped sensory spot; sdgco1, subdorsal type 1 glandular cell outlet; sdmgco2, subdorsal modified type 2 glandular cell outlet; sdrss, subdorsal rounded sensory spot; slne, sublateral nephridiopore; sls, sublateral spine; te, tergal extension; tsp, trichoscalid plate; vldss, ventrolateral droplet-shaped sensory spot; vmdss, ventromedial droplet-shaped sensory spot; vmgco1, ventromedial type 1 glandular cell outlet; vmrss, ventromedial rounded sensory spot." figureDoi="http://doi.org/10.5281/zenodo.10375299" httpUri="https://zenodo.org/record/10375299/files/figure.png" pageId="4" pageNumber="183">Fig. 3B</figureCitation>
;
<figureCitation id="BE9B2A1AC671FF8324EFF9247A90F93D" box="[655,686,1771,1790]" captionStart="Fig" captionStartId="7.[995,1025,149,166]" captionTargetBox="[102,968,150,1247]" captionTargetId="figure-378@7.[100,970,148,1249]" captionTargetPageId="7" captionText="Fig. 7. Scanning electron micrographs of additional specimens of Echinoderes angelae sp. nov. showing trunk segments and details of cuticular appendages. A: Subdorsal and laterodorsal sensory spots of segment 1, with the enlarged external ring of micropapillar (inset shows the intraspecific variability of these structures without such kind of external ring); B: ventral view of segments 2–6; C: subdorsal (top) and laterodorsal (bottom) modified type 2 glandular cell outlets of segments 2 and 4 respectively; D: laterodorsal view on left side of tergal plate of segment 2; E: nephridiopore and midlateral dropletshaped sensory spot of segment 9; F: subdorsal to midlateral view on right side of tergal plates of segments 8–9; G: middorsal spine of segment 4; H: subdorsal to midlateral view on left side of tergal plates of segments 3–4; I: laterodorsal tube of segment 10; J: lateral accessory tube of segment 8 (left) and sublateral spine of segment 7 (right); K: subdorsal type 1 glandular cell outlet of segment 9; L: ventral view of segments 10–11; M: subdorsal droplet-shaped sensory spot of segment 9. Abbreviations: go, gonopore; ms, muscular scar; spf, secondary pectinate fringe; modified type 2 glandular cell outlets are marked with continuous circles, and sensory spots with dashed teardrop-like figures." figureDoi="http://doi.org/10.5281/zenodo.10375309" httpUri="https://zenodo.org/record/10375309/files/figure.png" pageId="4" pageNumber="183">7G</figureCitation>
).
<typeStatus id="F91B883DC671FF8324A0F9247AD0F93D" box="[704,750,1771,1790]" pageId="4" pageNumber="183">Type</typeStatus>
1 glandular cell outlets in subdorsal and ventromedial positions (
<figureCitation id="BE9B2A1AC671FF8324DEF8C878A1F8F5" captionStart="Fig" captionStartId="3.[994,1024,149,166]" captionTargetBox="[101,967,149,1199]" captionTargetId="figure-599@3.[100,969,148,1201]" captionTargetPageId="3" captionText="Fig. 3. Line art drawing of Echinoderes angelae sp. nov. A: Ventral female overview; B: dorsal female overview; C: ventral male overview of segments 10–11; D: dorsal male overview of segments 10–11. Abbreviations: dpl, dorsal placid; go, gonopore; lat, lateral accessory tube; lddss, laterodorsal droplet-shaped sensory spot; ldmgco2, laterodorsal modified type 2 glandular cell outlet; ldrss, laterodorsal rounded sensory spot; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; lvgco1, lateroventral type 1 glandular cell outlet; lvmgco2, lateroventral modified type 2 glandular cell outlet; lvt, lateroventral tube; mddss, middorsal droplet-shaped sensory spot; mdgco1, middorsal type 1 glandular cell outlet; mds, middorsal spine; mldss, midlateral droplet-shaped sensory spot; mlmgco2, midlateral modified type 2 glandular cell outlet; ms, muscular scar; mvpl, midventral placid; ppf, primary pectinate fringe; ps, penile spines; S, segment (followed by number of corresponding segment); sddss, subdorsal droplet-shaped sensory spot; sdgco1, subdorsal type 1 glandular cell outlet; sdmgco2, subdorsal modified type 2 glandular cell outlet; sdrss, subdorsal rounded sensory spot; slne, sublateral nephridiopore; sls, sublateral spine; te, tergal extension; tsp, trichoscalid plate; vldss, ventrolateral droplet-shaped sensory spot; vmdss, ventromedial droplet-shaped sensory spot; vmgco1, ventromedial type 1 glandular cell outlet; vmrss, ventromedial rounded sensory spot." figureDoi="http://doi.org/10.5281/zenodo.10375299" httpUri="https://zenodo.org/record/10375299/files/figure.png" pageId="4" pageNumber="183">Fig. 3A and B</figureCitation>
). Modified
<typeStatus id="F91B883DC671FF83276EF8EC7909F8F5" box="[270,311,1827,1846]" pageId="4" pageNumber="183">type</typeStatus>
2 glandular cell outlets in laterodorsal position (
<figureCitation id="BE9B2A1AC671FF83260CF8F1788FF891" box="[108,177,1854,1874]" captionStart="Fig" captionStartId="3.[994,1024,149,166]" captionTargetBox="[101,967,149,1199]" captionTargetId="figure-599@3.[100,969,148,1201]" captionTargetPageId="3" captionText="Fig. 3. Line art drawing of Echinoderes angelae sp. nov. A: Ventral female overview; B: dorsal female overview; C: ventral male overview of segments 10–11; D: dorsal male overview of segments 10–11. Abbreviations: dpl, dorsal placid; go, gonopore; lat, lateral accessory tube; lddss, laterodorsal droplet-shaped sensory spot; ldmgco2, laterodorsal modified type 2 glandular cell outlet; ldrss, laterodorsal rounded sensory spot; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; lvgco1, lateroventral type 1 glandular cell outlet; lvmgco2, lateroventral modified type 2 glandular cell outlet; lvt, lateroventral tube; mddss, middorsal droplet-shaped sensory spot; mdgco1, middorsal type 1 glandular cell outlet; mds, middorsal spine; mldss, midlateral droplet-shaped sensory spot; mlmgco2, midlateral modified type 2 glandular cell outlet; ms, muscular scar; mvpl, midventral placid; ppf, primary pectinate fringe; ps, penile spines; S, segment (followed by number of corresponding segment); sddss, subdorsal droplet-shaped sensory spot; sdgco1, subdorsal type 1 glandular cell outlet; sdmgco2, subdorsal modified type 2 glandular cell outlet; sdrss, subdorsal rounded sensory spot; slne, sublateral nephridiopore; sls, sublateral spine; te, tergal extension; tsp, trichoscalid plate; vldss, ventrolateral droplet-shaped sensory spot; vmdss, ventromedial droplet-shaped sensory spot; vmgco1, ventromedial type 1 glandular cell outlet; vmrss, ventromedial rounded sensory spot." figureDoi="http://doi.org/10.5281/zenodo.10375299" httpUri="https://zenodo.org/record/10375299/files/figure.png" pageId="4" pageNumber="183">Fig. 3B</figureCitation>
;
<figureCitation id="BE9B2A1AC671FF8326DBF8F178E6F892" box="[187,216,1854,1873]" captionStart="Fig" captionStartId="7.[995,1025,149,166]" captionTargetBox="[102,968,150,1247]" captionTargetId="figure-378@7.[100,970,148,1249]" captionTargetPageId="7" captionText="Fig. 7. Scanning electron micrographs of additional specimens of Echinoderes angelae sp. nov. showing trunk segments and details of cuticular appendages. A: Subdorsal and laterodorsal sensory spots of segment 1, with the enlarged external ring of micropapillar (inset shows the intraspecific variability of these structures without such kind of external ring); B: ventral view of segments 2–6; C: subdorsal (top) and laterodorsal (bottom) modified type 2 glandular cell outlets of segments 2 and 4 respectively; D: laterodorsal view on left side of tergal plate of segment 2; E: nephridiopore and midlateral dropletshaped sensory spot of segment 9; F: subdorsal to midlateral view on right side of tergal plates of segments 8–9; G: middorsal spine of segment 4; H: subdorsal to midlateral view on left side of tergal plates of segments 3–4; I: laterodorsal tube of segment 10; J: lateral accessory tube of segment 8 (left) and sublateral spine of segment 7 (right); K: subdorsal type 1 glandular cell outlet of segment 9; L: ventral view of segments 10–11; M: subdorsal droplet-shaped sensory spot of segment 9. Abbreviations: go, gonopore; ms, muscular scar; spf, secondary pectinate fringe; modified type 2 glandular cell outlets are marked with continuous circles, and sensory spots with dashed teardrop-like figures." figureDoi="http://doi.org/10.5281/zenodo.10375309" httpUri="https://zenodo.org/record/10375309/files/figure.png" pageId="4" pageNumber="183">7C</figureCitation>
).
</paragraph>
<paragraph id="261F369FC671FF8326E4F8957DEEFBBF" blockId="4.[100,771,1073,1957]" lastBlockId="4.[818,1488,1073,1957]" pageId="4" pageNumber="183">
Segment 5 with tubes in lateroventral position (
<figureCitation id="BE9B2A1AC671FF832425F8957AB5F8AE" box="[581,651,1882,1901]" captionStart="Fig" captionStartId="3.[994,1024,149,166]" captionTargetBox="[101,967,149,1199]" captionTargetId="figure-599@3.[100,969,148,1201]" captionTargetPageId="3" captionText="Fig. 3. Line art drawing of Echinoderes angelae sp. nov. A: Ventral female overview; B: dorsal female overview; C: ventral male overview of segments 10–11; D: dorsal male overview of segments 10–11. Abbreviations: dpl, dorsal placid; go, gonopore; lat, lateral accessory tube; lddss, laterodorsal droplet-shaped sensory spot; ldmgco2, laterodorsal modified type 2 glandular cell outlet; ldrss, laterodorsal rounded sensory spot; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; lvgco1, lateroventral type 1 glandular cell outlet; lvmgco2, lateroventral modified type 2 glandular cell outlet; lvt, lateroventral tube; mddss, middorsal droplet-shaped sensory spot; mdgco1, middorsal type 1 glandular cell outlet; mds, middorsal spine; mldss, midlateral droplet-shaped sensory spot; mlmgco2, midlateral modified type 2 glandular cell outlet; ms, muscular scar; mvpl, midventral placid; ppf, primary pectinate fringe; ps, penile spines; S, segment (followed by number of corresponding segment); sddss, subdorsal droplet-shaped sensory spot; sdgco1, subdorsal type 1 glandular cell outlet; sdmgco2, subdorsal modified type 2 glandular cell outlet; sdrss, subdorsal rounded sensory spot; slne, sublateral nephridiopore; sls, sublateral spine; te, tergal extension; tsp, trichoscalid plate; vldss, ventrolateral droplet-shaped sensory spot; vmdss, ventromedial droplet-shaped sensory spot; vmgco1, ventromedial type 1 glandular cell outlet; vmrss, ventromedial rounded sensory spot." figureDoi="http://doi.org/10.5281/zenodo.10375299" httpUri="https://zenodo.org/record/10375299/files/figure.png" pageId="4" pageNumber="183">Fig. 3A</figureCitation>
;
<figureCitation id="BE9B2A1AC671FF8324F5F8957A8FF8AE" box="[661,689,1882,1901]" captionStart="Fig" captionStartId="5.[994,1024,149,166]" captionTargetBox="[102,967,150,1427]" captionTargetId="figure-287@5.[100,969,148,1429]" captionTargetPageId="5" captionText="Fig. 5. Light micrographs of female holotype MNHN- 655Ma (E), male paratype MNHN-661Ma (C–D), and additional specimens (A-B, F) of Echinoderes angelae sp. nov. showing trunk cuticular details. A: Middorsal to midlateral view on right side of tergal plate of segments 1–2; B: midlateral to ventromedial view on left side of cuticular plates of segments 5–7; C: ventral view of segments 5–6; D: dorsal view of segments 8–9; E: midlateral to ventromedial view on right side of cuticular plates of segments 9–10 (inset shows the intraspecific variability of the nephridiopore); F: midlateral to ventromedial view on right side of cuticular plates of segments 7–8. Abbreviations: lat, lateral accessory tube; lvt, lateroventral tube; ms, muscular scar; ppf, primary pectinate fringe; slne, sublateral nephridiopore; sls, sublateral spine; numbers after abbreviation indicate corresponding segment; glandular cell outlets are marked with continuous circles, and sensory spots with dashed circles or teardrop-like figures." figureDoi="http://doi.org/10.5281/zenodo.10375305" httpUri="https://zenodo.org/record/10375305/files/figure.png" pageId="4" pageNumber="183">5C</figureCitation>
).
<typeStatus id="F91B883DC671FF8324A2F8947ACEF8AD" box="[706,752,1883,1902]" pageId="4" pageNumber="183">Type</typeStatus>
1 glandular cell outlets in subdorsal and ventromedial positions (
<figureCitation id="BE9B2A1AC671FF8324DEF8B9789FF866" captionStart="Fig" captionStartId="3.[994,1024,149,166]" captionTargetBox="[101,967,149,1199]" captionTargetId="figure-599@3.[100,969,148,1201]" captionTargetPageId="3" captionText="Fig. 3. Line art drawing of Echinoderes angelae sp. nov. A: Ventral female overview; B: dorsal female overview; C: ventral male overview of segments 10–11; D: dorsal male overview of segments 10–11. Abbreviations: dpl, dorsal placid; go, gonopore; lat, lateral accessory tube; lddss, laterodorsal droplet-shaped sensory spot; ldmgco2, laterodorsal modified type 2 glandular cell outlet; ldrss, laterodorsal rounded sensory spot; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; lvgco1, lateroventral type 1 glandular cell outlet; lvmgco2, lateroventral modified type 2 glandular cell outlet; lvt, lateroventral tube; mddss, middorsal droplet-shaped sensory spot; mdgco1, middorsal type 1 glandular cell outlet; mds, middorsal spine; mldss, midlateral droplet-shaped sensory spot; mlmgco2, midlateral modified type 2 glandular cell outlet; ms, muscular scar; mvpl, midventral placid; ppf, primary pectinate fringe; ps, penile spines; S, segment (followed by number of corresponding segment); sddss, subdorsal droplet-shaped sensory spot; sdgco1, subdorsal type 1 glandular cell outlet; sdmgco2, subdorsal modified type 2 glandular cell outlet; sdrss, subdorsal rounded sensory spot; slne, sublateral nephridiopore; sls, sublateral spine; te, tergal extension; tsp, trichoscalid plate; vldss, ventrolateral droplet-shaped sensory spot; vmdss, ventromedial droplet-shaped sensory spot; vmgco1, ventromedial type 1 glandular cell outlet; vmrss, ventromedial rounded sensory spot." figureDoi="http://doi.org/10.5281/zenodo.10375299" httpUri="https://zenodo.org/record/10375299/files/figure.png" pageId="4" pageNumber="183">Fig. 3A and B</figureCitation>
). Modified
<typeStatus id="F91B883DC671FF832775F85D7900F866" box="[277,318,1938,1957]" pageId="4" pageNumber="183">type</typeStatus>
2 glandular cell outlets in subdorsal position (
<figureCitation id="BE9B2A1AC671FF83255AFBFE7BBDFB87" box="[826,899,1073,1092]" captionStart="Fig" captionStartId="3.[994,1024,149,166]" captionTargetBox="[101,967,149,1199]" captionTargetId="figure-599@3.[100,969,148,1201]" captionTargetPageId="3" captionText="Fig. 3. Line art drawing of Echinoderes angelae sp. nov. A: Ventral female overview; B: dorsal female overview; C: ventral male overview of segments 10–11; D: dorsal male overview of segments 10–11. Abbreviations: dpl, dorsal placid; go, gonopore; lat, lateral accessory tube; lddss, laterodorsal droplet-shaped sensory spot; ldmgco2, laterodorsal modified type 2 glandular cell outlet; ldrss, laterodorsal rounded sensory spot; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; lvgco1, lateroventral type 1 glandular cell outlet; lvmgco2, lateroventral modified type 2 glandular cell outlet; lvt, lateroventral tube; mddss, middorsal droplet-shaped sensory spot; mdgco1, middorsal type 1 glandular cell outlet; mds, middorsal spine; mldss, midlateral droplet-shaped sensory spot; mlmgco2, midlateral modified type 2 glandular cell outlet; ms, muscular scar; mvpl, midventral placid; ppf, primary pectinate fringe; ps, penile spines; S, segment (followed by number of corresponding segment); sddss, subdorsal droplet-shaped sensory spot; sdgco1, subdorsal type 1 glandular cell outlet; sdmgco2, subdorsal modified type 2 glandular cell outlet; sdrss, subdorsal rounded sensory spot; slne, sublateral nephridiopore; sls, sublateral spine; te, tergal extension; tsp, trichoscalid plate; vldss, ventrolateral droplet-shaped sensory spot; vmdss, ventromedial droplet-shaped sensory spot; vmgco1, ventromedial type 1 glandular cell outlet; vmrss, ventromedial rounded sensory spot." figureDoi="http://doi.org/10.5281/zenodo.10375299" httpUri="https://zenodo.org/record/10375299/files/figure.png" pageId="4" pageNumber="183">Fig. 3B</figureCitation>
). Droplet-shaped sensory spots in subdorsal, midlateral and ventromedial positions; subdorsal sensory spots more lateral than subdorsal modified
<typeStatus id="F91B883DC671FF8325ABFBA67BCAFBBF" box="[971,1012,1129,1148]" pageId="4" pageNumber="183">type</typeStatus>
2 glandular cell outlets (
<figureCitation id="BE9B2A1AC671FF832280FBA67D5CFBBF" box="[1248,1378,1129,1148]" captionStart="Fig" captionStartId="3.[994,1024,149,166]" captionTargetBox="[101,967,149,1199]" captionTargetId="figure-599@3.[100,969,148,1201]" captionTargetPageId="3" captionText="Fig. 3. Line art drawing of Echinoderes angelae sp. nov. A: Ventral female overview; B: dorsal female overview; C: ventral male overview of segments 10–11; D: dorsal male overview of segments 10–11. Abbreviations: dpl, dorsal placid; go, gonopore; lat, lateral accessory tube; lddss, laterodorsal droplet-shaped sensory spot; ldmgco2, laterodorsal modified type 2 glandular cell outlet; ldrss, laterodorsal rounded sensory spot; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; lvgco1, lateroventral type 1 glandular cell outlet; lvmgco2, lateroventral modified type 2 glandular cell outlet; lvt, lateroventral tube; mddss, middorsal droplet-shaped sensory spot; mdgco1, middorsal type 1 glandular cell outlet; mds, middorsal spine; mldss, midlateral droplet-shaped sensory spot; mlmgco2, midlateral modified type 2 glandular cell outlet; ms, muscular scar; mvpl, midventral placid; ppf, primary pectinate fringe; ps, penile spines; S, segment (followed by number of corresponding segment); sddss, subdorsal droplet-shaped sensory spot; sdgco1, subdorsal type 1 glandular cell outlet; sdmgco2, subdorsal modified type 2 glandular cell outlet; sdrss, subdorsal rounded sensory spot; slne, sublateral nephridiopore; sls, sublateral spine; te, tergal extension; tsp, trichoscalid plate; vldss, ventrolateral droplet-shaped sensory spot; vmdss, ventromedial droplet-shaped sensory spot; vmgco1, ventromedial type 1 glandular cell outlet; vmrss, ventromedial rounded sensory spot." figureDoi="http://doi.org/10.5281/zenodo.10375299" httpUri="https://zenodo.org/record/10375299/files/figure.png" pageId="4" pageNumber="183">Fig. 3A and B</figureCitation>
;
<figureCitation id="BE9B2A1AC671FF83230CFBA67DA1FBBF" box="[1388,1439,1129,1148]" captionStart="Fig" captionStartId="5.[994,1024,149,166]" captionTargetBox="[102,967,150,1427]" captionTargetId="figure-287@5.[100,969,148,1429]" captionTargetPageId="5" captionText="Fig. 5. Light micrographs of female holotype MNHN- 655Ma (E), male paratype MNHN-661Ma (C–D), and additional specimens (A-B, F) of Echinoderes angelae sp. nov. showing trunk cuticular details. A: Middorsal to midlateral view on right side of tergal plate of segments 1–2; B: midlateral to ventromedial view on left side of cuticular plates of segments 5–7; C: ventral view of segments 5–6; D: dorsal view of segments 8–9; E: midlateral to ventromedial view on right side of cuticular plates of segments 9–10 (inset shows the intraspecific variability of the nephridiopore); F: midlateral to ventromedial view on right side of cuticular plates of segments 7–8. Abbreviations: lat, lateral accessory tube; lvt, lateroventral tube; ms, muscular scar; ppf, primary pectinate fringe; slne, sublateral nephridiopore; sls, sublateral spine; numbers after abbreviation indicate corresponding segment; glandular cell outlets are marked with continuous circles, and sensory spots with dashed circles or teardrop-like figures." figureDoi="http://doi.org/10.5281/zenodo.10375305" httpUri="https://zenodo.org/record/10375305/files/figure.png" pageId="4" pageNumber="183">5B–C</figureCitation>
;
<figureCitation id="BE9B2A1AC671FF8323C8FBA67DFAFBBF" box="[1448,1476,1129,1148]" captionStart="Fig" captionStartId="7.[995,1025,149,166]" captionTargetBox="[102,968,150,1247]" captionTargetId="figure-378@7.[100,970,148,1249]" captionTargetPageId="7" captionText="Fig. 7. Scanning electron micrographs of additional specimens of Echinoderes angelae sp. nov. showing trunk segments and details of cuticular appendages. A: Subdorsal and laterodorsal sensory spots of segment 1, with the enlarged external ring of micropapillar (inset shows the intraspecific variability of these structures without such kind of external ring); B: ventral view of segments 2–6; C: subdorsal (top) and laterodorsal (bottom) modified type 2 glandular cell outlets of segments 2 and 4 respectively; D: laterodorsal view on left side of tergal plate of segment 2; E: nephridiopore and midlateral dropletshaped sensory spot of segment 9; F: subdorsal to midlateral view on right side of tergal plates of segments 8–9; G: middorsal spine of segment 4; H: subdorsal to midlateral view on left side of tergal plates of segments 3–4; I: laterodorsal tube of segment 10; J: lateral accessory tube of segment 8 (left) and sublateral spine of segment 7 (right); K: subdorsal type 1 glandular cell outlet of segment 9; L: ventral view of segments 10–11; M: subdorsal droplet-shaped sensory spot of segment 9. Abbreviations: go, gonopore; ms, muscular scar; spf, secondary pectinate fringe; modified type 2 glandular cell outlets are marked with continuous circles, and sensory spots with dashed teardrop-like figures." figureDoi="http://doi.org/10.5281/zenodo.10375309" httpUri="https://zenodo.org/record/10375309/files/figure.png" pageId="4" pageNumber="183">7B</figureCitation>
).
</paragraph>
<paragraph id="261F369FC671FF832531FB4A7C15FA98" blockId="4.[818,1488,1073,1957]" pageId="4" pageNumber="183">
Segment 6 with short (ca. 4–6 μm length), poorly sclerotized, weakly articulated, acicular spines in sublateral position (
<figureCitation id="BE9B2A1AC671FF83236EFB6E7D6BFB77" box="[1294,1365,1185,1204]" captionStart="Fig" captionStartId="3.[994,1024,149,166]" captionTargetBox="[101,967,149,1199]" captionTargetId="figure-599@3.[100,969,148,1201]" captionTargetPageId="3" captionText="Fig. 3. Line art drawing of Echinoderes angelae sp. nov. A: Ventral female overview; B: dorsal female overview; C: ventral male overview of segments 10–11; D: dorsal male overview of segments 10–11. Abbreviations: dpl, dorsal placid; go, gonopore; lat, lateral accessory tube; lddss, laterodorsal droplet-shaped sensory spot; ldmgco2, laterodorsal modified type 2 glandular cell outlet; ldrss, laterodorsal rounded sensory spot; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; lvgco1, lateroventral type 1 glandular cell outlet; lvmgco2, lateroventral modified type 2 glandular cell outlet; lvt, lateroventral tube; mddss, middorsal droplet-shaped sensory spot; mdgco1, middorsal type 1 glandular cell outlet; mds, middorsal spine; mldss, midlateral droplet-shaped sensory spot; mlmgco2, midlateral modified type 2 glandular cell outlet; ms, muscular scar; mvpl, midventral placid; ppf, primary pectinate fringe; ps, penile spines; S, segment (followed by number of corresponding segment); sddss, subdorsal droplet-shaped sensory spot; sdgco1, subdorsal type 1 glandular cell outlet; sdmgco2, subdorsal modified type 2 glandular cell outlet; sdrss, subdorsal rounded sensory spot; slne, sublateral nephridiopore; sls, sublateral spine; te, tergal extension; tsp, trichoscalid plate; vldss, ventrolateral droplet-shaped sensory spot; vmdss, ventromedial droplet-shaped sensory spot; vmgco1, ventromedial type 1 glandular cell outlet; vmrss, ventromedial rounded sensory spot." figureDoi="http://doi.org/10.5281/zenodo.10375299" httpUri="https://zenodo.org/record/10375299/files/figure.png" pageId="4" pageNumber="183">Fig. 3A</figureCitation>
;
<figureCitation id="BE9B2A1AC671FF832300FB6E7D42FB77" box="[1376,1404,1185,1204]" captionStart="Fig" captionStartId="5.[994,1024,149,166]" captionTargetBox="[102,967,150,1427]" captionTargetId="figure-287@5.[100,969,148,1429]" captionTargetPageId="5" captionText="Fig. 5. Light micrographs of female holotype MNHN- 655Ma (E), male paratype MNHN-661Ma (C–D), and additional specimens (A-B, F) of Echinoderes angelae sp. nov. showing trunk cuticular details. A: Middorsal to midlateral view on right side of tergal plate of segments 1–2; B: midlateral to ventromedial view on left side of cuticular plates of segments 5–7; C: ventral view of segments 5–6; D: dorsal view of segments 8–9; E: midlateral to ventromedial view on right side of cuticular plates of segments 9–10 (inset shows the intraspecific variability of the nephridiopore); F: midlateral to ventromedial view on right side of cuticular plates of segments 7–8. Abbreviations: lat, lateral accessory tube; lvt, lateroventral tube; ms, muscular scar; ppf, primary pectinate fringe; slne, sublateral nephridiopore; sls, sublateral spine; numbers after abbreviation indicate corresponding segment; glandular cell outlets are marked with continuous circles, and sensory spots with dashed circles or teardrop-like figures." figureDoi="http://doi.org/10.5281/zenodo.10375305" httpUri="https://zenodo.org/record/10375305/files/figure.png" pageId="4" pageNumber="183">5C</figureCitation>
).
<typeStatus id="F91B883DC671FF8323EEFB6E7D82FB77" box="[1422,1468,1185,1204]" pageId="4" pageNumber="183">Type</typeStatus>
1 glandular cell outlets in subdorsal and ventromedial positions (
<figureCitation id="BE9B2A1AC671FF8323EBFB727B53FB2F" captionStart="Fig" captionStartId="3.[994,1024,149,166]" captionTargetBox="[101,967,149,1199]" captionTargetId="figure-599@3.[100,969,148,1201]" captionTargetPageId="3" captionText="Fig. 3. Line art drawing of Echinoderes angelae sp. nov. A: Ventral female overview; B: dorsal female overview; C: ventral male overview of segments 10–11; D: dorsal male overview of segments 10–11. Abbreviations: dpl, dorsal placid; go, gonopore; lat, lateral accessory tube; lddss, laterodorsal droplet-shaped sensory spot; ldmgco2, laterodorsal modified type 2 glandular cell outlet; ldrss, laterodorsal rounded sensory spot; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; lvgco1, lateroventral type 1 glandular cell outlet; lvmgco2, lateroventral modified type 2 glandular cell outlet; lvt, lateroventral tube; mddss, middorsal droplet-shaped sensory spot; mdgco1, middorsal type 1 glandular cell outlet; mds, middorsal spine; mldss, midlateral droplet-shaped sensory spot; mlmgco2, midlateral modified type 2 glandular cell outlet; ms, muscular scar; mvpl, midventral placid; ppf, primary pectinate fringe; ps, penile spines; S, segment (followed by number of corresponding segment); sddss, subdorsal droplet-shaped sensory spot; sdgco1, subdorsal type 1 glandular cell outlet; sdmgco2, subdorsal modified type 2 glandular cell outlet; sdrss, subdorsal rounded sensory spot; slne, sublateral nephridiopore; sls, sublateral spine; te, tergal extension; tsp, trichoscalid plate; vldss, ventrolateral droplet-shaped sensory spot; vmdss, ventromedial droplet-shaped sensory spot; vmgco1, ventromedial type 1 glandular cell outlet; vmrss, ventromedial rounded sensory spot." figureDoi="http://doi.org/10.5281/zenodo.10375299" httpUri="https://zenodo.org/record/10375299/files/figure.png" pageId="4" pageNumber="183">Fig. 3A and B</figureCitation>
). Modified
<typeStatus id="F91B883DC671FF8325BEFB167C39FB2F" box="[990,1031,1241,1260]" pageId="4" pageNumber="183">type</typeStatus>
2 glandular cell outlets in subdorsal and midlateral positions (
<figureCitation id="BE9B2A1AC671FF8325BFFB3B7C19FACB" box="[991,1063,1268,1288]" captionStart="Fig" captionStartId="3.[994,1024,149,166]" captionTargetBox="[101,967,149,1199]" captionTargetId="figure-599@3.[100,969,148,1201]" captionTargetPageId="3" captionText="Fig. 3. Line art drawing of Echinoderes angelae sp. nov. A: Ventral female overview; B: dorsal female overview; C: ventral male overview of segments 10–11; D: dorsal male overview of segments 10–11. Abbreviations: dpl, dorsal placid; go, gonopore; lat, lateral accessory tube; lddss, laterodorsal droplet-shaped sensory spot; ldmgco2, laterodorsal modified type 2 glandular cell outlet; ldrss, laterodorsal rounded sensory spot; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; lvgco1, lateroventral type 1 glandular cell outlet; lvmgco2, lateroventral modified type 2 glandular cell outlet; lvt, lateroventral tube; mddss, middorsal droplet-shaped sensory spot; mdgco1, middorsal type 1 glandular cell outlet; mds, middorsal spine; mldss, midlateral droplet-shaped sensory spot; mlmgco2, midlateral modified type 2 glandular cell outlet; ms, muscular scar; mvpl, midventral placid; ppf, primary pectinate fringe; ps, penile spines; S, segment (followed by number of corresponding segment); sddss, subdorsal droplet-shaped sensory spot; sdgco1, subdorsal type 1 glandular cell outlet; sdmgco2, subdorsal modified type 2 glandular cell outlet; sdrss, subdorsal rounded sensory spot; slne, sublateral nephridiopore; sls, sublateral spine; te, tergal extension; tsp, trichoscalid plate; vldss, ventrolateral droplet-shaped sensory spot; vmdss, ventromedial droplet-shaped sensory spot; vmgco1, ventromedial type 1 glandular cell outlet; vmrss, ventromedial rounded sensory spot." figureDoi="http://doi.org/10.5281/zenodo.10375299" httpUri="https://zenodo.org/record/10375299/files/figure.png" pageId="4" pageNumber="183">Fig. 3B</figureCitation>
;
<figureCitation id="BE9B2A1AC671FF832255FB3B7C6CFAC4" box="[1077,1106,1268,1287]" captionStart="Fig" captionStartId="5.[994,1024,149,166]" captionTargetBox="[102,967,150,1427]" captionTargetId="figure-287@5.[100,969,148,1429]" captionTargetPageId="5" captionText="Fig. 5. Light micrographs of female holotype MNHN- 655Ma (E), male paratype MNHN-661Ma (C–D), and additional specimens (A-B, F) of Echinoderes angelae sp. nov. showing trunk cuticular details. A: Middorsal to midlateral view on right side of tergal plate of segments 1–2; B: midlateral to ventromedial view on left side of cuticular plates of segments 5–7; C: ventral view of segments 5–6; D: dorsal view of segments 8–9; E: midlateral to ventromedial view on right side of cuticular plates of segments 9–10 (inset shows the intraspecific variability of the nephridiopore); F: midlateral to ventromedial view on right side of cuticular plates of segments 7–8. Abbreviations: lat, lateral accessory tube; lvt, lateroventral tube; ms, muscular scar; ppf, primary pectinate fringe; slne, sublateral nephridiopore; sls, sublateral spine; numbers after abbreviation indicate corresponding segment; glandular cell outlets are marked with continuous circles, and sensory spots with dashed circles or teardrop-like figures." figureDoi="http://doi.org/10.5281/zenodo.10375305" httpUri="https://zenodo.org/record/10375305/files/figure.png" pageId="4" pageNumber="183">5B</figureCitation>
). Droplet-shaped sensory spots in subdorsal, midlateral and ventromedial positions; subdorsal sensory spots more mesial than subdorsal modified
<typeStatus id="F91B883DC671FF8322D1FAE27CE4FA83" box="[1201,1242,1325,1344]" pageId="4" pageNumber="183">type</typeStatus>
2 glandular cell outlets (
<figureCitation id="BE9B2A1AC671FF83255AFA877B82FA9F" box="[826,956,1352,1372]" captionStart="Fig" captionStartId="3.[994,1024,149,166]" captionTargetBox="[101,967,149,1199]" captionTargetId="figure-599@3.[100,969,148,1201]" captionTargetPageId="3" captionText="Fig. 3. Line art drawing of Echinoderes angelae sp. nov. A: Ventral female overview; B: dorsal female overview; C: ventral male overview of segments 10–11; D: dorsal male overview of segments 10–11. Abbreviations: dpl, dorsal placid; go, gonopore; lat, lateral accessory tube; lddss, laterodorsal droplet-shaped sensory spot; ldmgco2, laterodorsal modified type 2 glandular cell outlet; ldrss, laterodorsal rounded sensory spot; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; lvgco1, lateroventral type 1 glandular cell outlet; lvmgco2, lateroventral modified type 2 glandular cell outlet; lvt, lateroventral tube; mddss, middorsal droplet-shaped sensory spot; mdgco1, middorsal type 1 glandular cell outlet; mds, middorsal spine; mldss, midlateral droplet-shaped sensory spot; mlmgco2, midlateral modified type 2 glandular cell outlet; ms, muscular scar; mvpl, midventral placid; ppf, primary pectinate fringe; ps, penile spines; S, segment (followed by number of corresponding segment); sddss, subdorsal droplet-shaped sensory spot; sdgco1, subdorsal type 1 glandular cell outlet; sdmgco2, subdorsal modified type 2 glandular cell outlet; sdrss, subdorsal rounded sensory spot; slne, sublateral nephridiopore; sls, sublateral spine; te, tergal extension; tsp, trichoscalid plate; vldss, ventrolateral droplet-shaped sensory spot; vmdss, ventromedial droplet-shaped sensory spot; vmgco1, ventromedial type 1 glandular cell outlet; vmrss, ventromedial rounded sensory spot." figureDoi="http://doi.org/10.5281/zenodo.10375299" httpUri="https://zenodo.org/record/10375299/files/figure.png" pageId="4" pageNumber="183">Fig. 3A and B</figureCitation>
;
<figureCitation id="BE9B2A1AC671FF8325A6FA877BC4FA98" box="[966,1018,1352,1372]" captionStart="Fig" captionStartId="5.[994,1024,149,166]" captionTargetBox="[102,967,150,1427]" captionTargetId="figure-287@5.[100,969,148,1429]" captionTargetPageId="5" captionText="Fig. 5. Light micrographs of female holotype MNHN- 655Ma (E), male paratype MNHN-661Ma (C–D), and additional specimens (A-B, F) of Echinoderes angelae sp. nov. showing trunk cuticular details. A: Middorsal to midlateral view on right side of tergal plate of segments 1–2; B: midlateral to ventromedial view on left side of cuticular plates of segments 5–7; C: ventral view of segments 5–6; D: dorsal view of segments 8–9; E: midlateral to ventromedial view on right side of cuticular plates of segments 9–10 (inset shows the intraspecific variability of the nephridiopore); F: midlateral to ventromedial view on right side of cuticular plates of segments 7–8. Abbreviations: lat, lateral accessory tube; lvt, lateroventral tube; ms, muscular scar; ppf, primary pectinate fringe; slne, sublateral nephridiopore; sls, sublateral spine; numbers after abbreviation indicate corresponding segment; glandular cell outlets are marked with continuous circles, and sensory spots with dashed circles or teardrop-like figures." figureDoi="http://doi.org/10.5281/zenodo.10375305" httpUri="https://zenodo.org/record/10375305/files/figure.png" pageId="4" pageNumber="183">5B–C</figureCitation>
;
<figureCitation id="BE9B2A1AC671FF832263FA877C21FA98" box="[1027,1055,1352,1371]" captionStart="Fig" captionStartId="7.[995,1025,149,166]" captionTargetBox="[102,968,150,1247]" captionTargetId="figure-378@7.[100,970,148,1249]" captionTargetPageId="7" captionText="Fig. 7. Scanning electron micrographs of additional specimens of Echinoderes angelae sp. nov. showing trunk segments and details of cuticular appendages. A: Subdorsal and laterodorsal sensory spots of segment 1, with the enlarged external ring of micropapillar (inset shows the intraspecific variability of these structures without such kind of external ring); B: ventral view of segments 2–6; C: subdorsal (top) and laterodorsal (bottom) modified type 2 glandular cell outlets of segments 2 and 4 respectively; D: laterodorsal view on left side of tergal plate of segment 2; E: nephridiopore and midlateral dropletshaped sensory spot of segment 9; F: subdorsal to midlateral view on right side of tergal plates of segments 8–9; G: middorsal spine of segment 4; H: subdorsal to midlateral view on left side of tergal plates of segments 3–4; I: laterodorsal tube of segment 10; J: lateral accessory tube of segment 8 (left) and sublateral spine of segment 7 (right); K: subdorsal type 1 glandular cell outlet of segment 9; L: ventral view of segments 10–11; M: subdorsal droplet-shaped sensory spot of segment 9. Abbreviations: go, gonopore; ms, muscular scar; spf, secondary pectinate fringe; modified type 2 glandular cell outlets are marked with continuous circles, and sensory spots with dashed teardrop-like figures." figureDoi="http://doi.org/10.5281/zenodo.10375309" httpUri="https://zenodo.org/record/10375309/files/figure.png" pageId="4" pageNumber="183">7B</figureCitation>
).
</paragraph>
<paragraph id="261F369FC671FF832531FAAB7B68FA24" blockId="4.[818,1488,1073,1957]" pageId="4" pageNumber="183">
Segment 7 similar to segment 6 regarding spine, glandular cell outlet and sensory spot arrangement, but with modified
<typeStatus id="F91B883DC671FF83236CFA4F7D0BFA50" box="[1292,1333,1408,1427]" pageId="4" pageNumber="183">type</typeStatus>
2 glandular cell outlets in lateroventral instead of midlateral position and sensory spots in ventrolateral instead of ventromedial position (
<figureCitation id="BE9B2A1AC671FF83236BFA777DB0FA08" box="[1291,1422,1464,1483]" captionStart="Fig" captionStartId="3.[994,1024,149,166]" captionTargetBox="[101,967,149,1199]" captionTargetId="figure-599@3.[100,969,148,1201]" captionTargetPageId="3" captionText="Fig. 3. Line art drawing of Echinoderes angelae sp. nov. A: Ventral female overview; B: dorsal female overview; C: ventral male overview of segments 10–11; D: dorsal male overview of segments 10–11. Abbreviations: dpl, dorsal placid; go, gonopore; lat, lateral accessory tube; lddss, laterodorsal droplet-shaped sensory spot; ldmgco2, laterodorsal modified type 2 glandular cell outlet; ldrss, laterodorsal rounded sensory spot; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; lvgco1, lateroventral type 1 glandular cell outlet; lvmgco2, lateroventral modified type 2 glandular cell outlet; lvt, lateroventral tube; mddss, middorsal droplet-shaped sensory spot; mdgco1, middorsal type 1 glandular cell outlet; mds, middorsal spine; mldss, midlateral droplet-shaped sensory spot; mlmgco2, midlateral modified type 2 glandular cell outlet; ms, muscular scar; mvpl, midventral placid; ppf, primary pectinate fringe; ps, penile spines; S, segment (followed by number of corresponding segment); sddss, subdorsal droplet-shaped sensory spot; sdgco1, subdorsal type 1 glandular cell outlet; sdmgco2, subdorsal modified type 2 glandular cell outlet; sdrss, subdorsal rounded sensory spot; slne, sublateral nephridiopore; sls, sublateral spine; te, tergal extension; tsp, trichoscalid plate; vldss, ventrolateral droplet-shaped sensory spot; vmdss, ventromedial droplet-shaped sensory spot; vmgco1, ventromedial type 1 glandular cell outlet; vmrss, ventromedial rounded sensory spot." figureDoi="http://doi.org/10.5281/zenodo.10375299" httpUri="https://zenodo.org/record/10375299/files/figure.png" pageId="4" pageNumber="183">Fig. 3A and B</figureCitation>
;
<figureCitation id="BE9B2A1AC671FF8323F8FA777DF2FA08" box="[1432,1484,1464,1483]" captionStart="Fig" captionStartId="5.[994,1024,149,166]" captionTargetBox="[102,967,150,1427]" captionTargetId="figure-287@5.[100,969,148,1429]" captionTargetPageId="5" captionText="Fig. 5. Light micrographs of female holotype MNHN- 655Ma (E), male paratype MNHN-661Ma (C–D), and additional specimens (A-B, F) of Echinoderes angelae sp. nov. showing trunk cuticular details. A: Middorsal to midlateral view on right side of tergal plate of segments 1–2; B: midlateral to ventromedial view on left side of cuticular plates of segments 5–7; C: ventral view of segments 5–6; D: dorsal view of segments 8–9; E: midlateral to ventromedial view on right side of cuticular plates of segments 9–10 (inset shows the intraspecific variability of the nephridiopore); F: midlateral to ventromedial view on right side of cuticular plates of segments 7–8. Abbreviations: lat, lateral accessory tube; lvt, lateroventral tube; ms, muscular scar; ppf, primary pectinate fringe; slne, sublateral nephridiopore; sls, sublateral spine; numbers after abbreviation indicate corresponding segment; glandular cell outlets are marked with continuous circles, and sensory spots with dashed circles or teardrop-like figures." figureDoi="http://doi.org/10.5281/zenodo.10375305" httpUri="https://zenodo.org/record/10375305/files/figure.png" pageId="4" pageNumber="183">5B, F</figureCitation>
;
<figureCitation id="BE9B2A1AC671FF832552FA1B7B74FA24" box="[818,842,1492,1511]" captionStart="Fig" captionStartId="7.[995,1025,149,166]" captionTargetBox="[102,968,150,1247]" captionTargetId="figure-378@7.[100,970,148,1249]" captionTargetPageId="7" captionText="Fig. 7. Scanning electron micrographs of additional specimens of Echinoderes angelae sp. nov. showing trunk segments and details of cuticular appendages. A: Subdorsal and laterodorsal sensory spots of segment 1, with the enlarged external ring of micropapillar (inset shows the intraspecific variability of these structures without such kind of external ring); B: ventral view of segments 2–6; C: subdorsal (top) and laterodorsal (bottom) modified type 2 glandular cell outlets of segments 2 and 4 respectively; D: laterodorsal view on left side of tergal plate of segment 2; E: nephridiopore and midlateral dropletshaped sensory spot of segment 9; F: subdorsal to midlateral view on right side of tergal plates of segments 8–9; G: middorsal spine of segment 4; H: subdorsal to midlateral view on left side of tergal plates of segments 3–4; I: laterodorsal tube of segment 10; J: lateral accessory tube of segment 8 (left) and sublateral spine of segment 7 (right); K: subdorsal type 1 glandular cell outlet of segment 9; L: ventral view of segments 10–11; M: subdorsal droplet-shaped sensory spot of segment 9. Abbreviations: go, gonopore; ms, muscular scar; spf, secondary pectinate fringe; modified type 2 glandular cell outlets are marked with continuous circles, and sensory spots with dashed teardrop-like figures." figureDoi="http://doi.org/10.5281/zenodo.10375309" httpUri="https://zenodo.org/record/10375309/files/figure.png" pageId="4" pageNumber="183">7J</figureCitation>
).
</paragraph>
<paragraph id="261F369FC671FF832531FA3F7D87F9B1" blockId="4.[818,1488,1073,1957]" pageId="4" pageNumber="183">
Segment 8 with tubes in lateral accessory position (
<figureCitation id="BE9B2A1AC671FF832358FA207D40F9C0" box="[1336,1406,1519,1539]" captionStart="Fig" captionStartId="3.[994,1024,149,166]" captionTargetBox="[101,967,149,1199]" captionTargetId="figure-599@3.[100,969,148,1201]" captionTargetPageId="3" captionText="Fig. 3. Line art drawing of Echinoderes angelae sp. nov. A: Ventral female overview; B: dorsal female overview; C: ventral male overview of segments 10–11; D: dorsal male overview of segments 10–11. Abbreviations: dpl, dorsal placid; go, gonopore; lat, lateral accessory tube; lddss, laterodorsal droplet-shaped sensory spot; ldmgco2, laterodorsal modified type 2 glandular cell outlet; ldrss, laterodorsal rounded sensory spot; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; lvgco1, lateroventral type 1 glandular cell outlet; lvmgco2, lateroventral modified type 2 glandular cell outlet; lvt, lateroventral tube; mddss, middorsal droplet-shaped sensory spot; mdgco1, middorsal type 1 glandular cell outlet; mds, middorsal spine; mldss, midlateral droplet-shaped sensory spot; mlmgco2, midlateral modified type 2 glandular cell outlet; ms, muscular scar; mvpl, midventral placid; ppf, primary pectinate fringe; ps, penile spines; S, segment (followed by number of corresponding segment); sddss, subdorsal droplet-shaped sensory spot; sdgco1, subdorsal type 1 glandular cell outlet; sdmgco2, subdorsal modified type 2 glandular cell outlet; sdrss, subdorsal rounded sensory spot; slne, sublateral nephridiopore; sls, sublateral spine; te, tergal extension; tsp, trichoscalid plate; vldss, ventrolateral droplet-shaped sensory spot; vmdss, ventromedial droplet-shaped sensory spot; vmgco1, ventromedial type 1 glandular cell outlet; vmrss, ventromedial rounded sensory spot." figureDoi="http://doi.org/10.5281/zenodo.10375299" httpUri="https://zenodo.org/record/10375299/files/figure.png" pageId="4" pageNumber="183">Fig. 3A</figureCitation>
;
<figureCitation id="BE9B2A1AC671FF8323E7FA207D9FF9C1" box="[1415,1441,1519,1538]" captionStart="Fig" captionStartId="5.[994,1024,149,166]" captionTargetBox="[102,967,150,1427]" captionTargetId="figure-287@5.[100,969,148,1429]" captionTargetPageId="5" captionText="Fig. 5. Light micrographs of female holotype MNHN- 655Ma (E), male paratype MNHN-661Ma (C–D), and additional specimens (A-B, F) of Echinoderes angelae sp. nov. showing trunk cuticular details. A: Middorsal to midlateral view on right side of tergal plate of segments 1–2; B: midlateral to ventromedial view on left side of cuticular plates of segments 5–7; C: ventral view of segments 5–6; D: dorsal view of segments 8–9; E: midlateral to ventromedial view on right side of cuticular plates of segments 9–10 (inset shows the intraspecific variability of the nephridiopore); F: midlateral to ventromedial view on right side of cuticular plates of segments 7–8. Abbreviations: lat, lateral accessory tube; lvt, lateroventral tube; ms, muscular scar; ppf, primary pectinate fringe; slne, sublateral nephridiopore; sls, sublateral spine; numbers after abbreviation indicate corresponding segment; glandular cell outlets are marked with continuous circles, and sensory spots with dashed circles or teardrop-like figures." figureDoi="http://doi.org/10.5281/zenodo.10375305" httpUri="https://zenodo.org/record/10375305/files/figure.png" pageId="4" pageNumber="183">5F</figureCitation>
;
<figureCitation id="BE9B2A1AC671FF8323CBFA3F7DFDF9C0" box="[1451,1475,1520,1539]" captionStart="Fig" captionStartId="7.[995,1025,149,166]" captionTargetBox="[102,968,150,1247]" captionTargetId="figure-378@7.[100,970,148,1249]" captionTargetPageId="7" captionText="Fig. 7. Scanning electron micrographs of additional specimens of Echinoderes angelae sp. nov. showing trunk segments and details of cuticular appendages. A: Subdorsal and laterodorsal sensory spots of segment 1, with the enlarged external ring of micropapillar (inset shows the intraspecific variability of these structures without such kind of external ring); B: ventral view of segments 2–6; C: subdorsal (top) and laterodorsal (bottom) modified type 2 glandular cell outlets of segments 2 and 4 respectively; D: laterodorsal view on left side of tergal plate of segment 2; E: nephridiopore and midlateral dropletshaped sensory spot of segment 9; F: subdorsal to midlateral view on right side of tergal plates of segments 8–9; G: middorsal spine of segment 4; H: subdorsal to midlateral view on left side of tergal plates of segments 3–4; I: laterodorsal tube of segment 10; J: lateral accessory tube of segment 8 (left) and sublateral spine of segment 7 (right); K: subdorsal type 1 glandular cell outlet of segment 9; L: ventral view of segments 10–11; M: subdorsal droplet-shaped sensory spot of segment 9. Abbreviations: go, gonopore; ms, muscular scar; spf, secondary pectinate fringe; modified type 2 glandular cell outlets are marked with continuous circles, and sensory spots with dashed teardrop-like figures." figureDoi="http://doi.org/10.5281/zenodo.10375309" httpUri="https://zenodo.org/record/10375309/files/figure.png" pageId="4" pageNumber="183">7J</figureCitation>
).
<typeStatus id="F91B883DC671FF832552F9C37B5EF9DC" box="[818,864,1548,1567]" pageId="4" pageNumber="183">Type</typeStatus>
1 glandular cell outlets in subdorsal and ventromedial positions (
<figureCitation id="BE9B2A1AC671FF83255AF9E87BFBF9F9" box="[826,965,1575,1594]" captionStart="Fig" captionStartId="3.[994,1024,149,166]" captionTargetBox="[101,967,149,1199]" captionTargetId="figure-599@3.[100,969,148,1201]" captionTargetPageId="3" captionText="Fig. 3. Line art drawing of Echinoderes angelae sp. nov. A: Ventral female overview; B: dorsal female overview; C: ventral male overview of segments 10–11; D: dorsal male overview of segments 10–11. Abbreviations: dpl, dorsal placid; go, gonopore; lat, lateral accessory tube; lddss, laterodorsal droplet-shaped sensory spot; ldmgco2, laterodorsal modified type 2 glandular cell outlet; ldrss, laterodorsal rounded sensory spot; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; lvgco1, lateroventral type 1 glandular cell outlet; lvmgco2, lateroventral modified type 2 glandular cell outlet; lvt, lateroventral tube; mddss, middorsal droplet-shaped sensory spot; mdgco1, middorsal type 1 glandular cell outlet; mds, middorsal spine; mldss, midlateral droplet-shaped sensory spot; mlmgco2, midlateral modified type 2 glandular cell outlet; ms, muscular scar; mvpl, midventral placid; ppf, primary pectinate fringe; ps, penile spines; S, segment (followed by number of corresponding segment); sddss, subdorsal droplet-shaped sensory spot; sdgco1, subdorsal type 1 glandular cell outlet; sdmgco2, subdorsal modified type 2 glandular cell outlet; sdrss, subdorsal rounded sensory spot; slne, sublateral nephridiopore; sls, sublateral spine; te, tergal extension; tsp, trichoscalid plate; vldss, ventrolateral droplet-shaped sensory spot; vmdss, ventromedial droplet-shaped sensory spot; vmgco1, ventromedial type 1 glandular cell outlet; vmrss, ventromedial rounded sensory spot." figureDoi="http://doi.org/10.5281/zenodo.10375299" httpUri="https://zenodo.org/record/10375299/files/figure.png" pageId="4" pageNumber="183">Fig. 3A and B</figureCitation>
). Modified
<typeStatus id="F91B883DC671FF832257F9E77C5EF9F8" box="[1079,1120,1576,1595]" pageId="4" pageNumber="183">type</typeStatus>
2 glandular cell outlets in subdorsal, midlateral and lateroventral positions (
<figureCitation id="BE9B2A1AC671FF8322DBF98C7D71F994" box="[1211,1359,1603,1623]" captionStart="Fig" captionStartId="3.[994,1024,149,166]" captionTargetBox="[101,967,149,1199]" captionTargetId="figure-599@3.[100,969,148,1201]" captionTargetPageId="3" captionText="Fig. 3. Line art drawing of Echinoderes angelae sp. nov. A: Ventral female overview; B: dorsal female overview; C: ventral male overview of segments 10–11; D: dorsal male overview of segments 10–11. Abbreviations: dpl, dorsal placid; go, gonopore; lat, lateral accessory tube; lddss, laterodorsal droplet-shaped sensory spot; ldmgco2, laterodorsal modified type 2 glandular cell outlet; ldrss, laterodorsal rounded sensory spot; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; lvgco1, lateroventral type 1 glandular cell outlet; lvmgco2, lateroventral modified type 2 glandular cell outlet; lvt, lateroventral tube; mddss, middorsal droplet-shaped sensory spot; mdgco1, middorsal type 1 glandular cell outlet; mds, middorsal spine; mldss, midlateral droplet-shaped sensory spot; mlmgco2, midlateral modified type 2 glandular cell outlet; ms, muscular scar; mvpl, midventral placid; ppf, primary pectinate fringe; ps, penile spines; S, segment (followed by number of corresponding segment); sddss, subdorsal droplet-shaped sensory spot; sdgco1, subdorsal type 1 glandular cell outlet; sdmgco2, subdorsal modified type 2 glandular cell outlet; sdrss, subdorsal rounded sensory spot; slne, sublateral nephridiopore; sls, sublateral spine; te, tergal extension; tsp, trichoscalid plate; vldss, ventrolateral droplet-shaped sensory spot; vmdss, ventromedial droplet-shaped sensory spot; vmgco1, ventromedial type 1 glandular cell outlet; vmrss, ventromedial rounded sensory spot." figureDoi="http://doi.org/10.5281/zenodo.10375299" httpUri="https://zenodo.org/record/10375299/files/figure.png" pageId="4" pageNumber="183">Fig. 3A and B</figureCitation>
;
<figureCitation id="BE9B2A1AC671FF83233FF98C7DA4F994" box="[1375,1434,1603,1623]" captionStart="Fig" captionStartId="5.[994,1024,149,166]" captionTargetBox="[102,967,150,1427]" captionTargetId="figure-287@5.[100,969,148,1429]" captionTargetPageId="5" captionText="Fig. 5. Light micrographs of female holotype MNHN- 655Ma (E), male paratype MNHN-661Ma (C–D), and additional specimens (A-B, F) of Echinoderes angelae sp. nov. showing trunk cuticular details. A: Middorsal to midlateral view on right side of tergal plate of segments 1–2; B: midlateral to ventromedial view on left side of cuticular plates of segments 5–7; C: ventral view of segments 5–6; D: dorsal view of segments 8–9; E: midlateral to ventromedial view on right side of cuticular plates of segments 9–10 (inset shows the intraspecific variability of the nephridiopore); F: midlateral to ventromedial view on right side of cuticular plates of segments 7–8. Abbreviations: lat, lateral accessory tube; lvt, lateroventral tube; ms, muscular scar; ppf, primary pectinate fringe; slne, sublateral nephridiopore; sls, sublateral spine; numbers after abbreviation indicate corresponding segment; glandular cell outlets are marked with continuous circles, and sensory spots with dashed circles or teardrop-like figures." figureDoi="http://doi.org/10.5281/zenodo.10375305" httpUri="https://zenodo.org/record/10375305/files/figure.png" pageId="4" pageNumber="183">5D, F</figureCitation>
;
<figureCitation id="BE9B2A1AC671FF8323CAF98C7DFBF995" box="[1450,1477,1603,1622]" captionStart="Fig" captionStartId="7.[995,1025,149,166]" captionTargetBox="[102,968,150,1247]" captionTargetId="figure-378@7.[100,970,148,1249]" captionTargetPageId="7" captionText="Fig. 7. Scanning electron micrographs of additional specimens of Echinoderes angelae sp. nov. showing trunk segments and details of cuticular appendages. A: Subdorsal and laterodorsal sensory spots of segment 1, with the enlarged external ring of micropapillar (inset shows the intraspecific variability of these structures without such kind of external ring); B: ventral view of segments 2–6; C: subdorsal (top) and laterodorsal (bottom) modified type 2 glandular cell outlets of segments 2 and 4 respectively; D: laterodorsal view on left side of tergal plate of segment 2; E: nephridiopore and midlateral dropletshaped sensory spot of segment 9; F: subdorsal to midlateral view on right side of tergal plates of segments 8–9; G: middorsal spine of segment 4; H: subdorsal to midlateral view on left side of tergal plates of segments 3–4; I: laterodorsal tube of segment 10; J: lateral accessory tube of segment 8 (left) and sublateral spine of segment 7 (right); K: subdorsal type 1 glandular cell outlet of segment 9; L: ventral view of segments 10–11; M: subdorsal droplet-shaped sensory spot of segment 9. Abbreviations: go, gonopore; ms, muscular scar; spf, secondary pectinate fringe; modified type 2 glandular cell outlets are marked with continuous circles, and sensory spots with dashed teardrop-like figures." figureDoi="http://doi.org/10.5281/zenodo.10375309" httpUri="https://zenodo.org/record/10375309/files/figure.png" pageId="4" pageNumber="183">7F</figureCitation>
). Droplet-shaped sensory spots in subdorsal position (
<figureCitation id="BE9B2A1AC671FF83237DF9907D5FF9B1" box="[1309,1377,1631,1650]" captionStart="Fig" captionStartId="3.[994,1024,149,166]" captionTargetBox="[101,967,149,1199]" captionTargetId="figure-599@3.[100,969,148,1201]" captionTargetPageId="3" captionText="Fig. 3. Line art drawing of Echinoderes angelae sp. nov. A: Ventral female overview; B: dorsal female overview; C: ventral male overview of segments 10–11; D: dorsal male overview of segments 10–11. Abbreviations: dpl, dorsal placid; go, gonopore; lat, lateral accessory tube; lddss, laterodorsal droplet-shaped sensory spot; ldmgco2, laterodorsal modified type 2 glandular cell outlet; ldrss, laterodorsal rounded sensory spot; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; lvgco1, lateroventral type 1 glandular cell outlet; lvmgco2, lateroventral modified type 2 glandular cell outlet; lvt, lateroventral tube; mddss, middorsal droplet-shaped sensory spot; mdgco1, middorsal type 1 glandular cell outlet; mds, middorsal spine; mldss, midlateral droplet-shaped sensory spot; mlmgco2, midlateral modified type 2 glandular cell outlet; ms, muscular scar; mvpl, midventral placid; ppf, primary pectinate fringe; ps, penile spines; S, segment (followed by number of corresponding segment); sddss, subdorsal droplet-shaped sensory spot; sdgco1, subdorsal type 1 glandular cell outlet; sdmgco2, subdorsal modified type 2 glandular cell outlet; sdrss, subdorsal rounded sensory spot; slne, sublateral nephridiopore; sls, sublateral spine; te, tergal extension; tsp, trichoscalid plate; vldss, ventrolateral droplet-shaped sensory spot; vmdss, ventromedial droplet-shaped sensory spot; vmgco1, ventromedial type 1 glandular cell outlet; vmrss, ventromedial rounded sensory spot." figureDoi="http://doi.org/10.5281/zenodo.10375299" httpUri="https://zenodo.org/record/10375299/files/figure.png" pageId="4" pageNumber="183">Fig. 3B</figureCitation>
;
<figureCitation id="BE9B2A1AC671FF83230CF9907DB6F9B1" box="[1388,1416,1631,1650]" captionStart="Fig" captionStartId="5.[994,1024,149,166]" captionTargetBox="[102,967,150,1427]" captionTargetId="figure-287@5.[100,969,148,1429]" captionTargetPageId="5" captionText="Fig. 5. Light micrographs of female holotype MNHN- 655Ma (E), male paratype MNHN-661Ma (C–D), and additional specimens (A-B, F) of Echinoderes angelae sp. nov. showing trunk cuticular details. A: Middorsal to midlateral view on right side of tergal plate of segments 1–2; B: midlateral to ventromedial view on left side of cuticular plates of segments 5–7; C: ventral view of segments 5–6; D: dorsal view of segments 8–9; E: midlateral to ventromedial view on right side of cuticular plates of segments 9–10 (inset shows the intraspecific variability of the nephridiopore); F: midlateral to ventromedial view on right side of cuticular plates of segments 7–8. Abbreviations: lat, lateral accessory tube; lvt, lateroventral tube; ms, muscular scar; ppf, primary pectinate fringe; slne, sublateral nephridiopore; sls, sublateral spine; numbers after abbreviation indicate corresponding segment; glandular cell outlets are marked with continuous circles, and sensory spots with dashed circles or teardrop-like figures." figureDoi="http://doi.org/10.5281/zenodo.10375305" httpUri="https://zenodo.org/record/10375305/files/figure.png" pageId="4" pageNumber="183">5D</figureCitation>
;
<figureCitation id="BE9B2A1AC671FF8323F3F9907D90F9B1" box="[1427,1454,1631,1650]" captionStart="Fig" captionStartId="7.[995,1025,149,166]" captionTargetBox="[102,968,150,1247]" captionTargetId="figure-378@7.[100,970,148,1249]" captionTargetPageId="7" captionText="Fig. 7. Scanning electron micrographs of additional specimens of Echinoderes angelae sp. nov. showing trunk segments and details of cuticular appendages. A: Subdorsal and laterodorsal sensory spots of segment 1, with the enlarged external ring of micropapillar (inset shows the intraspecific variability of these structures without such kind of external ring); B: ventral view of segments 2–6; C: subdorsal (top) and laterodorsal (bottom) modified type 2 glandular cell outlets of segments 2 and 4 respectively; D: laterodorsal view on left side of tergal plate of segment 2; E: nephridiopore and midlateral dropletshaped sensory spot of segment 9; F: subdorsal to midlateral view on right side of tergal plates of segments 8–9; G: middorsal spine of segment 4; H: subdorsal to midlateral view on left side of tergal plates of segments 3–4; I: laterodorsal tube of segment 10; J: lateral accessory tube of segment 8 (left) and sublateral spine of segment 7 (right); K: subdorsal type 1 glandular cell outlet of segment 9; L: ventral view of segments 10–11; M: subdorsal droplet-shaped sensory spot of segment 9. Abbreviations: go, gonopore; ms, muscular scar; spf, secondary pectinate fringe; modified type 2 glandular cell outlets are marked with continuous circles, and sensory spots with dashed teardrop-like figures." figureDoi="http://doi.org/10.5281/zenodo.10375309" httpUri="https://zenodo.org/record/10375309/files/figure.png" pageId="4" pageNumber="183">7F</figureCitation>
).
</paragraph>
<paragraph id="261F369FC671FF832531F9B47D8DF8F5" blockId="4.[818,1488,1073,1957]" pageId="4" pageNumber="183">
Segment 9 with
<typeStatus id="F91B883DC671FF83259DF9B47C18F94D" box="[1021,1062,1659,1678]" pageId="4" pageNumber="183">type</typeStatus>
1 glandular cell outlets in subdorsal and ventromedial positions (
<figureCitation id="BE9B2A1AC671FF832277F9587CA4F969" box="[1047,1178,1687,1706]" captionStart="Fig" captionStartId="3.[994,1024,149,166]" captionTargetBox="[101,967,149,1199]" captionTargetId="figure-599@3.[100,969,148,1201]" captionTargetPageId="3" captionText="Fig. 3. Line art drawing of Echinoderes angelae sp. nov. A: Ventral female overview; B: dorsal female overview; C: ventral male overview of segments 10–11; D: dorsal male overview of segments 10–11. Abbreviations: dpl, dorsal placid; go, gonopore; lat, lateral accessory tube; lddss, laterodorsal droplet-shaped sensory spot; ldmgco2, laterodorsal modified type 2 glandular cell outlet; ldrss, laterodorsal rounded sensory spot; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; lvgco1, lateroventral type 1 glandular cell outlet; lvmgco2, lateroventral modified type 2 glandular cell outlet; lvt, lateroventral tube; mddss, middorsal droplet-shaped sensory spot; mdgco1, middorsal type 1 glandular cell outlet; mds, middorsal spine; mldss, midlateral droplet-shaped sensory spot; mlmgco2, midlateral modified type 2 glandular cell outlet; ms, muscular scar; mvpl, midventral placid; ppf, primary pectinate fringe; ps, penile spines; S, segment (followed by number of corresponding segment); sddss, subdorsal droplet-shaped sensory spot; sdgco1, subdorsal type 1 glandular cell outlet; sdmgco2, subdorsal modified type 2 glandular cell outlet; sdrss, subdorsal rounded sensory spot; slne, sublateral nephridiopore; sls, sublateral spine; te, tergal extension; tsp, trichoscalid plate; vldss, ventrolateral droplet-shaped sensory spot; vmdss, ventromedial droplet-shaped sensory spot; vmgco1, ventromedial type 1 glandular cell outlet; vmrss, ventromedial rounded sensory spot." figureDoi="http://doi.org/10.5281/zenodo.10375299" httpUri="https://zenodo.org/record/10375299/files/figure.png" pageId="4" pageNumber="183">Fig. 3A and B</figureCitation>
;
<figureCitation id="BE9B2A1AC671FF8322C4F9587CFFF969" box="[1188,1217,1687,1706]" captionStart="Fig" captionStartId="7.[995,1025,149,166]" captionTargetBox="[102,968,150,1247]" captionTargetId="figure-378@7.[100,970,148,1249]" captionTargetPageId="7" captionText="Fig. 7. Scanning electron micrographs of additional specimens of Echinoderes angelae sp. nov. showing trunk segments and details of cuticular appendages. A: Subdorsal and laterodorsal sensory spots of segment 1, with the enlarged external ring of micropapillar (inset shows the intraspecific variability of these structures without such kind of external ring); B: ventral view of segments 2–6; C: subdorsal (top) and laterodorsal (bottom) modified type 2 glandular cell outlets of segments 2 and 4 respectively; D: laterodorsal view on left side of tergal plate of segment 2; E: nephridiopore and midlateral dropletshaped sensory spot of segment 9; F: subdorsal to midlateral view on right side of tergal plates of segments 8–9; G: middorsal spine of segment 4; H: subdorsal to midlateral view on left side of tergal plates of segments 3–4; I: laterodorsal tube of segment 10; J: lateral accessory tube of segment 8 (left) and sublateral spine of segment 7 (right); K: subdorsal type 1 glandular cell outlet of segment 9; L: ventral view of segments 10–11; M: subdorsal droplet-shaped sensory spot of segment 9. Abbreviations: go, gonopore; ms, muscular scar; spf, secondary pectinate fringe; modified type 2 glandular cell outlets are marked with continuous circles, and sensory spots with dashed teardrop-like figures." figureDoi="http://doi.org/10.5281/zenodo.10375309" httpUri="https://zenodo.org/record/10375309/files/figure.png" pageId="4" pageNumber="183">7K</figureCitation>
). One pair of droplet-shaped sensory spots in midlateral and ventrolateral positions, and two pairs in subdorsal position (
<figureCitation id="BE9B2A1AC671FF832269F9007CB1F921" box="[1033,1167,1743,1762]" captionStart="Fig" captionStartId="3.[994,1024,149,166]" captionTargetBox="[101,967,149,1199]" captionTargetId="figure-599@3.[100,969,148,1201]" captionTargetPageId="3" captionText="Fig. 3. Line art drawing of Echinoderes angelae sp. nov. A: Ventral female overview; B: dorsal female overview; C: ventral male overview of segments 10–11; D: dorsal male overview of segments 10–11. Abbreviations: dpl, dorsal placid; go, gonopore; lat, lateral accessory tube; lddss, laterodorsal droplet-shaped sensory spot; ldmgco2, laterodorsal modified type 2 glandular cell outlet; ldrss, laterodorsal rounded sensory spot; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; lvgco1, lateroventral type 1 glandular cell outlet; lvmgco2, lateroventral modified type 2 glandular cell outlet; lvt, lateroventral tube; mddss, middorsal droplet-shaped sensory spot; mdgco1, middorsal type 1 glandular cell outlet; mds, middorsal spine; mldss, midlateral droplet-shaped sensory spot; mlmgco2, midlateral modified type 2 glandular cell outlet; ms, muscular scar; mvpl, midventral placid; ppf, primary pectinate fringe; ps, penile spines; S, segment (followed by number of corresponding segment); sddss, subdorsal droplet-shaped sensory spot; sdgco1, subdorsal type 1 glandular cell outlet; sdmgco2, subdorsal modified type 2 glandular cell outlet; sdrss, subdorsal rounded sensory spot; slne, sublateral nephridiopore; sls, sublateral spine; te, tergal extension; tsp, trichoscalid plate; vldss, ventrolateral droplet-shaped sensory spot; vmdss, ventromedial droplet-shaped sensory spot; vmgco1, ventromedial type 1 glandular cell outlet; vmrss, ventromedial rounded sensory spot." figureDoi="http://doi.org/10.5281/zenodo.10375299" httpUri="https://zenodo.org/record/10375299/files/figure.png" pageId="4" pageNumber="183">Fig. 3A and B</figureCitation>
;
<figureCitation id="BE9B2A1AC671FF8322FBF9007C9BF921" box="[1179,1189,1743,1762]" captionStart="Fig" captionStartId="5.[994,1024,149,166]" captionTargetBox="[102,967,150,1427]" captionTargetId="figure-287@5.[100,969,148,1429]" captionTargetPageId="5" captionText="Fig. 5. Light micrographs of female holotype MNHN- 655Ma (E), male paratype MNHN-661Ma (C–D), and additional specimens (A-B, F) of Echinoderes angelae sp. nov. showing trunk cuticular details. A: Middorsal to midlateral view on right side of tergal plate of segments 1–2; B: midlateral to ventromedial view on left side of cuticular plates of segments 5–7; C: ventral view of segments 5–6; D: dorsal view of segments 8–9; E: midlateral to ventromedial view on right side of cuticular plates of segments 9–10 (inset shows the intraspecific variability of the nephridiopore); F: midlateral to ventromedial view on right side of cuticular plates of segments 7–8. Abbreviations: lat, lateral accessory tube; lvt, lateroventral tube; ms, muscular scar; ppf, primary pectinate fringe; slne, sublateral nephridiopore; sls, sublateral spine; numbers after abbreviation indicate corresponding segment; glandular cell outlets are marked with continuous circles, and sensory spots with dashed circles or teardrop-like figures." figureDoi="http://doi.org/10.5281/zenodo.10375305" httpUri="https://zenodo.org/record/10375305/files/figure.png" pageId="4" pageNumber="183">5</figureCitation>
D-E; 7F, M). Nephridiopores in sublateral position as enlarged sieve plate openings with an anterior, elongated, slightly convex area with numerous pores and a posterior, slightly concave region with a single, central pore (
<figureCitation id="BE9B2A1AC671FF832377F8ED7D60F8F5" box="[1303,1374,1826,1846]" captionStart="Fig" captionStartId="3.[994,1024,149,166]" captionTargetBox="[101,967,149,1199]" captionTargetId="figure-599@3.[100,969,148,1201]" captionTargetPageId="3" captionText="Fig. 3. Line art drawing of Echinoderes angelae sp. nov. A: Ventral female overview; B: dorsal female overview; C: ventral male overview of segments 10–11; D: dorsal male overview of segments 10–11. Abbreviations: dpl, dorsal placid; go, gonopore; lat, lateral accessory tube; lddss, laterodorsal droplet-shaped sensory spot; ldmgco2, laterodorsal modified type 2 glandular cell outlet; ldrss, laterodorsal rounded sensory spot; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; lvgco1, lateroventral type 1 glandular cell outlet; lvmgco2, lateroventral modified type 2 glandular cell outlet; lvt, lateroventral tube; mddss, middorsal droplet-shaped sensory spot; mdgco1, middorsal type 1 glandular cell outlet; mds, middorsal spine; mldss, midlateral droplet-shaped sensory spot; mlmgco2, midlateral modified type 2 glandular cell outlet; ms, muscular scar; mvpl, midventral placid; ppf, primary pectinate fringe; ps, penile spines; S, segment (followed by number of corresponding segment); sddss, subdorsal droplet-shaped sensory spot; sdgco1, subdorsal type 1 glandular cell outlet; sdmgco2, subdorsal modified type 2 glandular cell outlet; sdrss, subdorsal rounded sensory spot; slne, sublateral nephridiopore; sls, sublateral spine; te, tergal extension; tsp, trichoscalid plate; vldss, ventrolateral droplet-shaped sensory spot; vmdss, ventromedial droplet-shaped sensory spot; vmgco1, ventromedial type 1 glandular cell outlet; vmrss, ventromedial rounded sensory spot." figureDoi="http://doi.org/10.5281/zenodo.10375299" httpUri="https://zenodo.org/record/10375299/files/figure.png" pageId="4" pageNumber="183">Fig. 3A</figureCitation>
;
<figureCitation id="BE9B2A1AC671FF832307F8ED7DBCF8F6" box="[1383,1410,1826,1845]" captionStart="Fig" captionStartId="5.[994,1024,149,166]" captionTargetBox="[102,967,150,1427]" captionTargetId="figure-287@5.[100,969,148,1429]" captionTargetPageId="5" captionText="Fig. 5. Light micrographs of female holotype MNHN- 655Ma (E), male paratype MNHN-661Ma (C–D), and additional specimens (A-B, F) of Echinoderes angelae sp. nov. showing trunk cuticular details. A: Middorsal to midlateral view on right side of tergal plate of segments 1–2; B: midlateral to ventromedial view on left side of cuticular plates of segments 5–7; C: ventral view of segments 5–6; D: dorsal view of segments 8–9; E: midlateral to ventromedial view on right side of cuticular plates of segments 9–10 (inset shows the intraspecific variability of the nephridiopore); F: midlateral to ventromedial view on right side of cuticular plates of segments 7–8. Abbreviations: lat, lateral accessory tube; lvt, lateroventral tube; ms, muscular scar; ppf, primary pectinate fringe; slne, sublateral nephridiopore; sls, sublateral spine; numbers after abbreviation indicate corresponding segment; glandular cell outlets are marked with continuous circles, and sensory spots with dashed circles or teardrop-like figures." figureDoi="http://doi.org/10.5281/zenodo.10375305" httpUri="https://zenodo.org/record/10375305/files/figure.png" pageId="4" pageNumber="183">5E</figureCitation>
;
<figureCitation id="BE9B2A1AC671FF8323ECF8EC7D96F8F5" box="[1420,1448,1827,1846]" captionStart="Fig" captionStartId="7.[995,1025,149,166]" captionTargetBox="[102,968,150,1247]" captionTargetId="figure-378@7.[100,970,148,1249]" captionTargetPageId="7" captionText="Fig. 7. Scanning electron micrographs of additional specimens of Echinoderes angelae sp. nov. showing trunk segments and details of cuticular appendages. A: Subdorsal and laterodorsal sensory spots of segment 1, with the enlarged external ring of micropapillar (inset shows the intraspecific variability of these structures without such kind of external ring); B: ventral view of segments 2–6; C: subdorsal (top) and laterodorsal (bottom) modified type 2 glandular cell outlets of segments 2 and 4 respectively; D: laterodorsal view on left side of tergal plate of segment 2; E: nephridiopore and midlateral dropletshaped sensory spot of segment 9; F: subdorsal to midlateral view on right side of tergal plates of segments 8–9; G: middorsal spine of segment 4; H: subdorsal to midlateral view on left side of tergal plates of segments 3–4; I: laterodorsal tube of segment 10; J: lateral accessory tube of segment 8 (left) and sublateral spine of segment 7 (right); K: subdorsal type 1 glandular cell outlet of segment 9; L: ventral view of segments 10–11; M: subdorsal droplet-shaped sensory spot of segment 9. Abbreviations: go, gonopore; ms, muscular scar; spf, secondary pectinate fringe; modified type 2 glandular cell outlets are marked with continuous circles, and sensory spots with dashed teardrop-like figures." figureDoi="http://doi.org/10.5281/zenodo.10375309" httpUri="https://zenodo.org/record/10375309/files/figure.png" pageId="4" pageNumber="183">7E</figureCitation>
).
</paragraph>
<paragraph id="261F369FC671FF822531F8F1791FF9F9" blockId="4.[818,1488,1073,1957]" lastBlockId="5.[100,770,1519,1873]" lastPageId="5" lastPageNumber="184" pageId="4" pageNumber="183">
Segment 10 with tubes in laterodorsal position, larger in males (
<figureCitation id="BE9B2A1AC671FF83255AF8957BA4F8AD" box="[826,922,1882,1902]" captionStart="Fig" captionStartId="3.[994,1024,149,166]" captionTargetBox="[101,967,149,1199]" captionTargetId="figure-599@3.[100,969,148,1201]" captionTargetPageId="3" captionText="Fig. 3. Line art drawing of Echinoderes angelae sp. nov. A: Ventral female overview; B: dorsal female overview; C: ventral male overview of segments 10–11; D: dorsal male overview of segments 10–11. Abbreviations: dpl, dorsal placid; go, gonopore; lat, lateral accessory tube; lddss, laterodorsal droplet-shaped sensory spot; ldmgco2, laterodorsal modified type 2 glandular cell outlet; ldrss, laterodorsal rounded sensory spot; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; lvgco1, lateroventral type 1 glandular cell outlet; lvmgco2, lateroventral modified type 2 glandular cell outlet; lvt, lateroventral tube; mddss, middorsal droplet-shaped sensory spot; mdgco1, middorsal type 1 glandular cell outlet; mds, middorsal spine; mldss, midlateral droplet-shaped sensory spot; mlmgco2, midlateral modified type 2 glandular cell outlet; ms, muscular scar; mvpl, midventral placid; ppf, primary pectinate fringe; ps, penile spines; S, segment (followed by number of corresponding segment); sddss, subdorsal droplet-shaped sensory spot; sdgco1, subdorsal type 1 glandular cell outlet; sdmgco2, subdorsal modified type 2 glandular cell outlet; sdrss, subdorsal rounded sensory spot; slne, sublateral nephridiopore; sls, sublateral spine; te, tergal extension; tsp, trichoscalid plate; vldss, ventrolateral droplet-shaped sensory spot; vmdss, ventromedial droplet-shaped sensory spot; vmgco1, ventromedial type 1 glandular cell outlet; vmrss, ventromedial rounded sensory spot." figureDoi="http://doi.org/10.5281/zenodo.10375299" httpUri="https://zenodo.org/record/10375299/files/figure.png" pageId="4" pageNumber="183">Fig. 3B, D</figureCitation>
;
<figureCitation id="BE9B2A1AC671FF8325C5F8957BFEF8AE" box="[933,960,1882,1901]" captionStart="Fig" captionStartId="4.[995,1025,149,166]" captionTargetBox="[102,968,150,977]" captionTargetId="figure-749@4.[100,970,148,979]" captionTargetPageId="4" captionText="Fig. 4. Light micrographs of female holotype MNHN- 655Ma (A, C-D), male paratype MNHN-661Ma (B, E), female paratypes MNHN-656Ma (H) and MNHN-657Ma (F), and additional specimens (G, I) of Echinoderes angelae sp. nov. showing trunk overview, details of mouth cone and neck, epibionts and ingested diatoms. A: Ventral trunk overview; B: mouth cone, showing the outer oral styles; C: dorsal neck view; D: ventral neck view; E: middorsal to laterodorsal view on right side of tergal plate of segments 10–11; F: lateral view of segment 11; G: ingested diatom; H: unknown type of epibionts attached to the posterior edge of segment 11; I: filamentous bacteria attached to the posterior edge of segment 11. Abbreviations: ba, bacteria; bs, basal sheath; dpl, dorsal placid; ep, end-piece; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; mvpl, midventral placid; ps, penile spines; te, tergal extension; tsp, trichoscalid plate." figureDoi="http://doi.org/10.5281/zenodo.10375301" httpUri="https://zenodo.org/record/10375301/files/figure.png" pageId="4" pageNumber="183">4E</figureCitation>
;
<figureCitation id="BE9B2A1AC671FF8325AAF8957BDEF8AE" box="[970,992,1882,1901]" captionStart="Fig" captionStartId="7.[995,1025,149,166]" captionTargetBox="[102,968,150,1247]" captionTargetId="figure-378@7.[100,970,148,1249]" captionTargetPageId="7" captionText="Fig. 7. Scanning electron micrographs of additional specimens of Echinoderes angelae sp. nov. showing trunk segments and details of cuticular appendages. A: Subdorsal and laterodorsal sensory spots of segment 1, with the enlarged external ring of micropapillar (inset shows the intraspecific variability of these structures without such kind of external ring); B: ventral view of segments 2–6; C: subdorsal (top) and laterodorsal (bottom) modified type 2 glandular cell outlets of segments 2 and 4 respectively; D: laterodorsal view on left side of tergal plate of segment 2; E: nephridiopore and midlateral dropletshaped sensory spot of segment 9; F: subdorsal to midlateral view on right side of tergal plates of segments 8–9; G: middorsal spine of segment 4; H: subdorsal to midlateral view on left side of tergal plates of segments 3–4; I: laterodorsal tube of segment 10; J: lateral accessory tube of segment 8 (left) and sublateral spine of segment 7 (right); K: subdorsal type 1 glandular cell outlet of segment 9; L: ventral view of segments 10–11; M: subdorsal droplet-shaped sensory spot of segment 9. Abbreviations: go, gonopore; ms, muscular scar; spf, secondary pectinate fringe; modified type 2 glandular cell outlets are marked with continuous circles, and sensory spots with dashed teardrop-like figures." figureDoi="http://doi.org/10.5281/zenodo.10375309" httpUri="https://zenodo.org/record/10375309/files/figure.png" pageId="4" pageNumber="183">7I</figureCitation>
).
<specimenCount id="30A6FD16C671FF832592F8947C75F8AD" box="[1010,1099,1883,1902]" count="2" pageId="4" pageNumber="183" type="generic" typeStatus="type">Two type</specimenCount>
1 glandular cell outlets in middorsal position, longitudinally arranged;
<typeStatus id="F91B883DC671FF832209F8B87CACF849" box="[1129,1170,1911,1930]" pageId="4" pageNumber="183">type</typeStatus>
1 glandular cell outlets also in ventromedial position (
<figureCitation id="BE9B2A1AC671FF832272F85D7CA7F866" box="[1042,1177,1938,1957]" captionStart="Fig" captionStartId="3.[994,1024,149,166]" captionTargetBox="[101,967,149,1199]" captionTargetId="figure-599@3.[100,969,148,1201]" captionTargetPageId="3" captionText="Fig. 3. Line art drawing of Echinoderes angelae sp. nov. A: Ventral female overview; B: dorsal female overview; C: ventral male overview of segments 10–11; D: dorsal male overview of segments 10–11. Abbreviations: dpl, dorsal placid; go, gonopore; lat, lateral accessory tube; lddss, laterodorsal droplet-shaped sensory spot; ldmgco2, laterodorsal modified type 2 glandular cell outlet; ldrss, laterodorsal rounded sensory spot; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; lvgco1, lateroventral type 1 glandular cell outlet; lvmgco2, lateroventral modified type 2 glandular cell outlet; lvt, lateroventral tube; mddss, middorsal droplet-shaped sensory spot; mdgco1, middorsal type 1 glandular cell outlet; mds, middorsal spine; mldss, midlateral droplet-shaped sensory spot; mlmgco2, midlateral modified type 2 glandular cell outlet; ms, muscular scar; mvpl, midventral placid; ppf, primary pectinate fringe; ps, penile spines; S, segment (followed by number of corresponding segment); sddss, subdorsal droplet-shaped sensory spot; sdgco1, subdorsal type 1 glandular cell outlet; sdmgco2, subdorsal modified type 2 glandular cell outlet; sdrss, subdorsal rounded sensory spot; slne, sublateral nephridiopore; sls, sublateral spine; te, tergal extension; tsp, trichoscalid plate; vldss, ventrolateral droplet-shaped sensory spot; vmdss, ventromedial droplet-shaped sensory spot; vmgco1, ventromedial type 1 glandular cell outlet; vmrss, ventromedial rounded sensory spot." figureDoi="http://doi.org/10.5281/zenodo.10375299" httpUri="https://zenodo.org/record/10375299/files/figure.png" pageId="4" pageNumber="183">Fig. 3A and B</figureCitation>
). Modified
<typeStatus id="F91B883DC671FF832368F85D7D0FF866" box="[1288,1329,1938,1957]" pageId="4" pageNumber="183">type</typeStatus>
2 glandular cell outlets in laterodorsal position, anterior to the tubes (
<figureCitation id="BE9B2A1AC670FF82243FFA207A9AF9C1" box="[607,676,1519,1538]" captionStart="Fig" captionStartId="3.[994,1024,149,166]" captionTargetBox="[101,967,149,1199]" captionTargetId="figure-599@3.[100,969,148,1201]" captionTargetPageId="3" captionText="Fig. 3. Line art drawing of Echinoderes angelae sp. nov. A: Ventral female overview; B: dorsal female overview; C: ventral male overview of segments 10–11; D: dorsal male overview of segments 10–11. Abbreviations: dpl, dorsal placid; go, gonopore; lat, lateral accessory tube; lddss, laterodorsal droplet-shaped sensory spot; ldmgco2, laterodorsal modified type 2 glandular cell outlet; ldrss, laterodorsal rounded sensory spot; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; lvgco1, lateroventral type 1 glandular cell outlet; lvmgco2, lateroventral modified type 2 glandular cell outlet; lvt, lateroventral tube; mddss, middorsal droplet-shaped sensory spot; mdgco1, middorsal type 1 glandular cell outlet; mds, middorsal spine; mldss, midlateral droplet-shaped sensory spot; mlmgco2, midlateral modified type 2 glandular cell outlet; ms, muscular scar; mvpl, midventral placid; ppf, primary pectinate fringe; ps, penile spines; S, segment (followed by number of corresponding segment); sddss, subdorsal droplet-shaped sensory spot; sdgco1, subdorsal type 1 glandular cell outlet; sdmgco2, subdorsal modified type 2 glandular cell outlet; sdrss, subdorsal rounded sensory spot; slne, sublateral nephridiopore; sls, sublateral spine; te, tergal extension; tsp, trichoscalid plate; vldss, ventrolateral droplet-shaped sensory spot; vmdss, ventromedial droplet-shaped sensory spot; vmgco1, ventromedial type 1 glandular cell outlet; vmrss, ventromedial rounded sensory spot." figureDoi="http://doi.org/10.5281/zenodo.10375299" httpUri="https://zenodo.org/record/10375299/files/figure.png" pageId="5" pageNumber="184">Fig. 3B</figureCitation>
). Droplet-shaped sensory spots in subdorsal and ventrolateral positions (
<figureCitation id="BE9B2A1AC670FF82260CF9E878F2F9F9" box="[108,204,1575,1594]" captionStart="Fig" captionStartId="3.[994,1024,149,166]" captionTargetBox="[101,967,149,1199]" captionTargetId="figure-599@3.[100,969,148,1201]" captionTargetPageId="3" captionText="Fig. 3. Line art drawing of Echinoderes angelae sp. nov. A: Ventral female overview; B: dorsal female overview; C: ventral male overview of segments 10–11; D: dorsal male overview of segments 10–11. Abbreviations: dpl, dorsal placid; go, gonopore; lat, lateral accessory tube; lddss, laterodorsal droplet-shaped sensory spot; ldmgco2, laterodorsal modified type 2 glandular cell outlet; ldrss, laterodorsal rounded sensory spot; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; lvgco1, lateroventral type 1 glandular cell outlet; lvmgco2, lateroventral modified type 2 glandular cell outlet; lvt, lateroventral tube; mddss, middorsal droplet-shaped sensory spot; mdgco1, middorsal type 1 glandular cell outlet; mds, middorsal spine; mldss, midlateral droplet-shaped sensory spot; mlmgco2, midlateral modified type 2 glandular cell outlet; ms, muscular scar; mvpl, midventral placid; ppf, primary pectinate fringe; ps, penile spines; S, segment (followed by number of corresponding segment); sddss, subdorsal droplet-shaped sensory spot; sdgco1, subdorsal type 1 glandular cell outlet; sdmgco2, subdorsal modified type 2 glandular cell outlet; sdrss, subdorsal rounded sensory spot; slne, sublateral nephridiopore; sls, sublateral spine; te, tergal extension; tsp, trichoscalid plate; vldss, ventrolateral droplet-shaped sensory spot; vmdss, ventromedial droplet-shaped sensory spot; vmgco1, ventromedial type 1 glandular cell outlet; vmrss, ventromedial rounded sensory spot." figureDoi="http://doi.org/10.5281/zenodo.10375299" httpUri="https://zenodo.org/record/10375299/files/figure.png" pageId="5" pageNumber="184">Fig. 3A–D</figureCitation>
;
<figureCitation id="BE9B2A1AC670FF8226B6F9E878CFF9F9" box="[214,241,1575,1594]" captionStart="Fig" captionStartId="5.[994,1024,149,166]" captionTargetBox="[102,967,150,1427]" captionTargetId="figure-287@5.[100,969,148,1429]" captionTargetPageId="5" captionText="Fig. 5. Light micrographs of female holotype MNHN- 655Ma (E), male paratype MNHN-661Ma (C–D), and additional specimens (A-B, F) of Echinoderes angelae sp. nov. showing trunk cuticular details. A: Middorsal to midlateral view on right side of tergal plate of segments 1–2; B: midlateral to ventromedial view on left side of cuticular plates of segments 5–7; C: ventral view of segments 5–6; D: dorsal view of segments 8–9; E: midlateral to ventromedial view on right side of cuticular plates of segments 9–10 (inset shows the intraspecific variability of the nephridiopore); F: midlateral to ventromedial view on right side of cuticular plates of segments 7–8. Abbreviations: lat, lateral accessory tube; lvt, lateroventral tube; ms, muscular scar; ppf, primary pectinate fringe; slne, sublateral nephridiopore; sls, sublateral spine; numbers after abbreviation indicate corresponding segment; glandular cell outlets are marked with continuous circles, and sensory spots with dashed circles or teardrop-like figures." figureDoi="http://doi.org/10.5281/zenodo.10375305" httpUri="https://zenodo.org/record/10375305/files/figure.png" pageId="5" pageNumber="184">5E</figureCitation>
;
<figureCitation id="BE9B2A1AC670FF82269AF9E87928F9F9" box="[250,278,1575,1594]" captionStart="Fig" captionStartId="7.[995,1025,149,166]" captionTargetBox="[102,968,150,1247]" captionTargetId="figure-378@7.[100,970,148,1249]" captionTargetPageId="7" captionText="Fig. 7. Scanning electron micrographs of additional specimens of Echinoderes angelae sp. nov. showing trunk segments and details of cuticular appendages. A: Subdorsal and laterodorsal sensory spots of segment 1, with the enlarged external ring of micropapillar (inset shows the intraspecific variability of these structures without such kind of external ring); B: ventral view of segments 2–6; C: subdorsal (top) and laterodorsal (bottom) modified type 2 glandular cell outlets of segments 2 and 4 respectively; D: laterodorsal view on left side of tergal plate of segment 2; E: nephridiopore and midlateral dropletshaped sensory spot of segment 9; F: subdorsal to midlateral view on right side of tergal plates of segments 8–9; G: middorsal spine of segment 4; H: subdorsal to midlateral view on left side of tergal plates of segments 3–4; I: laterodorsal tube of segment 10; J: lateral accessory tube of segment 8 (left) and sublateral spine of segment 7 (right); K: subdorsal type 1 glandular cell outlet of segment 9; L: ventral view of segments 10–11; M: subdorsal droplet-shaped sensory spot of segment 9. Abbreviations: go, gonopore; ms, muscular scar; spf, secondary pectinate fringe; modified type 2 glandular cell outlets are marked with continuous circles, and sensory spots with dashed teardrop-like figures." figureDoi="http://doi.org/10.5281/zenodo.10375309" httpUri="https://zenodo.org/record/10375309/files/figure.png" pageId="5" pageNumber="184">7L</figureCitation>
).
</paragraph>
<caption id="72DF6617C670FF822582FF5A7DEEFD9B" ID-DOI="http://doi.org/10.5281/zenodo.10375305" ID-Zenodo-Dep="10375305" httpUri="https://zenodo.org/record/10375305/files/figure.png" pageId="5" pageNumber="184" startId="5.[994,1024,149,166]" targetBox="[102,967,150,1427]" targetPageId="5" targetType="figure">
<paragraph id="261F369FC670FF822582FF5A7DEEFD9B" blockId="5.[994,1488,149,600]" pageId="5" pageNumber="184">
<emphasis id="14D4EA8DC670FF822582FF5A7C1FFF65" bold="true" box="[994,1057,149,166]" pageId="5" pageNumber="184">Fig. 5.</emphasis>
Light micrographs of female holotype MNHN- 655Ma (E), male paratype MNHN-661Ma (C–D), and additional specimens (A-B, F) of
<taxonomicName id="E1A04D1CC670FF82236AFF077D90FF1A" authority="Cepeda &amp; Gayet &amp; Spedicato &amp; Michaud &amp; Zeppilli, 2022" authorityName="Cepeda &amp; Gayet &amp; Spedicato &amp; Michaud &amp; Zeppilli" authorityYear="2022" box="[1290,1454,200,217]" class="Cyclorhagida" family="Echinoderidae" genus="Echinoderes" kingdom="Animalia" order="Echinorhagata" pageId="5" pageNumber="184" phylum="Kinorhyncha" rank="species" species="angelae" status="sp. nov.">
<emphasis id="14D4EA8DC670FF82236AFF077D90FF1A" bold="true" box="[1290,1454,200,217]" italics="true" pageId="5" pageNumber="184">Echinoderes angelae</emphasis>
</taxonomicName>
<taxonomicNameLabel id="0FE757F6C670FF8223D7FF067C36FF31" pageId="5" pageNumber="184" rank="species">sp. nov.</taxonomicNameLabel>
showing trunk cuticular details. A: Middorsal to midlateral view on right side of tergal plate of segments 1–2; B: midlateral to ventromedial view on left side of cuticular plates of segments 5–7; C: ventral view of segments 5–6; D: dorsal view of segments 8–9; E: midlateral to ventromedial view on right side of cuticular plates of segments 9–10 (inset shows the intraspecific variability of the nephridiopore); F: midlateral to ventromedial view on right side of cuticular plates of segments 7–8. Abbreviations: lat, lateral accessory tube; lvt, lateroventral tube; ms, muscular scar; ppf, primary pectinate fringe; slne, sublateral nephridiopore; sls, sublateral spine; numbers after abbreviation indicate corresponding segment; glandular cell outlets are marked with continuous circles, and sensory spots with dashed circles or teardrop-like figures.
</paragraph>
</caption>
<paragraph id="261F369FC670FF8226E4F98C7C92F9F9" blockId="5.[100,770,1519,1873]" lastBlockId="5.[818,1487,1519,1594]" pageId="5" pageNumber="184">
Segment 11 with relatively short lateral terminal spines (LTS:TL average ratio = 17.9%), distally pointed, with hollow central cavity, superficially covered by hairs (
<figureCitation id="BE9B2A1AC670FF8227E7F9B479D8F94D" box="[391,486,1659,1678]" captionStart="Fig" captionStartId="3.[994,1024,149,166]" captionTargetBox="[101,967,149,1199]" captionTargetId="figure-599@3.[100,969,148,1201]" captionTargetPageId="3" captionText="Fig. 3. Line art drawing of Echinoderes angelae sp. nov. A: Ventral female overview; B: dorsal female overview; C: ventral male overview of segments 10–11; D: dorsal male overview of segments 10–11. Abbreviations: dpl, dorsal placid; go, gonopore; lat, lateral accessory tube; lddss, laterodorsal droplet-shaped sensory spot; ldmgco2, laterodorsal modified type 2 glandular cell outlet; ldrss, laterodorsal rounded sensory spot; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; lvgco1, lateroventral type 1 glandular cell outlet; lvmgco2, lateroventral modified type 2 glandular cell outlet; lvt, lateroventral tube; mddss, middorsal droplet-shaped sensory spot; mdgco1, middorsal type 1 glandular cell outlet; mds, middorsal spine; mldss, midlateral droplet-shaped sensory spot; mlmgco2, midlateral modified type 2 glandular cell outlet; ms, muscular scar; mvpl, midventral placid; ppf, primary pectinate fringe; ps, penile spines; S, segment (followed by number of corresponding segment); sddss, subdorsal droplet-shaped sensory spot; sdgco1, subdorsal type 1 glandular cell outlet; sdmgco2, subdorsal modified type 2 glandular cell outlet; sdrss, subdorsal rounded sensory spot; slne, sublateral nephridiopore; sls, sublateral spine; te, tergal extension; tsp, trichoscalid plate; vldss, ventrolateral droplet-shaped sensory spot; vmdss, ventromedial droplet-shaped sensory spot; vmgco1, ventromedial type 1 glandular cell outlet; vmrss, ventromedial rounded sensory spot." figureDoi="http://doi.org/10.5281/zenodo.10375299" httpUri="https://zenodo.org/record/10375299/files/figure.png" pageId="5" pageNumber="184">Fig. 3A–D</figureCitation>
;
<figureCitation id="BE9B2A1AC670FF822790F9B47A33F94D" box="[496,525,1659,1678]" captionStart="Fig" captionStartId="4.[995,1025,149,166]" captionTargetBox="[102,968,150,977]" captionTargetId="figure-749@4.[100,970,148,979]" captionTargetPageId="4" captionText="Fig. 4. Light micrographs of female holotype MNHN- 655Ma (A, C-D), male paratype MNHN-661Ma (B, E), female paratypes MNHN-656Ma (H) and MNHN-657Ma (F), and additional specimens (G, I) of Echinoderes angelae sp. nov. showing trunk overview, details of mouth cone and neck, epibionts and ingested diatoms. A: Ventral trunk overview; B: mouth cone, showing the outer oral styles; C: dorsal neck view; D: ventral neck view; E: middorsal to laterodorsal view on right side of tergal plate of segments 10–11; F: lateral view of segment 11; G: ingested diatom; H: unknown type of epibionts attached to the posterior edge of segment 11; I: filamentous bacteria attached to the posterior edge of segment 11. Abbreviations: ba, bacteria; bs, basal sheath; dpl, dorsal placid; ep, end-piece; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; mvpl, midventral placid; ps, penile spines; te, tergal extension; tsp, trichoscalid plate." figureDoi="http://doi.org/10.5281/zenodo.10375301" httpUri="https://zenodo.org/record/10375301/files/figure.png" pageId="5" pageNumber="184">4A</figureCitation>
, E-F, H–I; 6A, E-G). Males with three pairs of penile spines that emerge from laterodorsal position in the intersegmental joint with the preceding segment; penile spines tube-like, covered by hairs, abruptly tapering near their distal tips (
<figureCitation id="BE9B2A1AC670FF82260CF925788FF93D" box="[108,177,1770,1790]" captionStart="Fig" captionStartId="3.[994,1024,149,166]" captionTargetBox="[101,967,149,1199]" captionTargetId="figure-599@3.[100,969,148,1201]" captionTargetPageId="3" captionText="Fig. 3. Line art drawing of Echinoderes angelae sp. nov. A: Ventral female overview; B: dorsal female overview; C: ventral male overview of segments 10–11; D: dorsal male overview of segments 10–11. Abbreviations: dpl, dorsal placid; go, gonopore; lat, lateral accessory tube; lddss, laterodorsal droplet-shaped sensory spot; ldmgco2, laterodorsal modified type 2 glandular cell outlet; ldrss, laterodorsal rounded sensory spot; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; lvgco1, lateroventral type 1 glandular cell outlet; lvmgco2, lateroventral modified type 2 glandular cell outlet; lvt, lateroventral tube; mddss, middorsal droplet-shaped sensory spot; mdgco1, middorsal type 1 glandular cell outlet; mds, middorsal spine; mldss, midlateral droplet-shaped sensory spot; mlmgco2, midlateral modified type 2 glandular cell outlet; ms, muscular scar; mvpl, midventral placid; ppf, primary pectinate fringe; ps, penile spines; S, segment (followed by number of corresponding segment); sddss, subdorsal droplet-shaped sensory spot; sdgco1, subdorsal type 1 glandular cell outlet; sdmgco2, subdorsal modified type 2 glandular cell outlet; sdrss, subdorsal rounded sensory spot; slne, sublateral nephridiopore; sls, sublateral spine; te, tergal extension; tsp, trichoscalid plate; vldss, ventrolateral droplet-shaped sensory spot; vmdss, ventromedial droplet-shaped sensory spot; vmgco1, ventromedial type 1 glandular cell outlet; vmrss, ventromedial rounded sensory spot." figureDoi="http://doi.org/10.5281/zenodo.10375299" httpUri="https://zenodo.org/record/10375299/files/figure.png" pageId="5" pageNumber="184">Fig. 3D</figureCitation>
;
<figureCitation id="BE9B2A1AC670FF8226DAF92578EBF93E" box="[186,213,1770,1789]" captionStart="Fig" captionStartId="4.[995,1025,149,166]" captionTargetBox="[102,968,150,977]" captionTargetId="figure-749@4.[100,970,148,979]" captionTargetPageId="4" captionText="Fig. 4. Light micrographs of female holotype MNHN- 655Ma (A, C-D), male paratype MNHN-661Ma (B, E), female paratypes MNHN-656Ma (H) and MNHN-657Ma (F), and additional specimens (G, I) of Echinoderes angelae sp. nov. showing trunk overview, details of mouth cone and neck, epibionts and ingested diatoms. A: Ventral trunk overview; B: mouth cone, showing the outer oral styles; C: dorsal neck view; D: ventral neck view; E: middorsal to laterodorsal view on right side of tergal plate of segments 10–11; F: lateral view of segment 11; G: ingested diatom; H: unknown type of epibionts attached to the posterior edge of segment 11; I: filamentous bacteria attached to the posterior edge of segment 11. Abbreviations: ba, bacteria; bs, basal sheath; dpl, dorsal placid; ep, end-piece; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; mvpl, midventral placid; ps, penile spines; te, tergal extension; tsp, trichoscalid plate." figureDoi="http://doi.org/10.5281/zenodo.10375301" httpUri="https://zenodo.org/record/10375301/files/figure.png" pageId="5" pageNumber="184">4E</figureCitation>
;
<figureCitation id="BE9B2A1AC670FF8226BDF92578C6F93E" box="[221,248,1770,1789]" captionStart="Fig" captionStartId="6.[997,1027,149,166]" captionTargetBox="[102,970,150,1260]" captionTargetId="figure-385@6.[100,972,148,1261]" captionTargetPageId="6" captionText="Fig. 6. Scanning electron micrographs of additional specimens of Echinoderes angelae sp. nov. showing trunk overview and details of introvert, neck, lateral terminal spines and sexually dimorphic features. A: Lateral trunk overview; B: introvert, sector 3; C: introvert, sector 4; D: ventral neck view; E: lateral terminal spine; F: male penile spines; G: female lateral terminal accessory spine. Abbreviations: ba, bacteria; ftz, fringed transverse zone; ltas, lateral terminal accessory spine; lts, lateral terminal spine; mvpl, midventral placid; ne, neck; ps, penile spines; psp, primary spinoscalid; sp, spinoscalid (followed by number of corresponding ring); te, tergal extension; ts, trichoscalid; tsp, trichoscalid plate; lambda marks (̂) indicate attachment point of spinoscalids." figureDoi="http://doi.org/10.5281/zenodo.10375307" httpUri="https://zenodo.org/record/10375307/files/figure.png" pageId="5" pageNumber="184">6F</figureCitation>
). Females with short (LTAS:LTS average ratio = 28.5%), distally frayed, covered by hairs, lateral terminal accessory spines, and oval, ventrolateral gonopores near the intersegmental joint with the preceding segment (
<figureCitation id="BE9B2A1AC670FF822740F8F179A2F892" box="[288,412,1854,1873]" captionStart="Fig" captionStartId="3.[994,1024,149,166]" captionTargetBox="[101,967,149,1199]" captionTargetId="figure-599@3.[100,969,148,1201]" captionTargetPageId="3" captionText="Fig. 3. Line art drawing of Echinoderes angelae sp. nov. A: Ventral female overview; B: dorsal female overview; C: ventral male overview of segments 10–11; D: dorsal male overview of segments 10–11. Abbreviations: dpl, dorsal placid; go, gonopore; lat, lateral accessory tube; lddss, laterodorsal droplet-shaped sensory spot; ldmgco2, laterodorsal modified type 2 glandular cell outlet; ldrss, laterodorsal rounded sensory spot; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; lvgco1, lateroventral type 1 glandular cell outlet; lvmgco2, lateroventral modified type 2 glandular cell outlet; lvt, lateroventral tube; mddss, middorsal droplet-shaped sensory spot; mdgco1, middorsal type 1 glandular cell outlet; mds, middorsal spine; mldss, midlateral droplet-shaped sensory spot; mlmgco2, midlateral modified type 2 glandular cell outlet; ms, muscular scar; mvpl, midventral placid; ppf, primary pectinate fringe; ps, penile spines; S, segment (followed by number of corresponding segment); sddss, subdorsal droplet-shaped sensory spot; sdgco1, subdorsal type 1 glandular cell outlet; sdmgco2, subdorsal modified type 2 glandular cell outlet; sdrss, subdorsal rounded sensory spot; slne, sublateral nephridiopore; sls, sublateral spine; te, tergal extension; tsp, trichoscalid plate; vldss, ventrolateral droplet-shaped sensory spot; vmdss, ventromedial droplet-shaped sensory spot; vmgco1, ventromedial type 1 glandular cell outlet; vmrss, ventromedial rounded sensory spot." figureDoi="http://doi.org/10.5281/zenodo.10375299" httpUri="https://zenodo.org/record/10375299/files/figure.png" pageId="5" pageNumber="184">Fig. 3A and B</figureCitation>
;
<figureCitation id="BE9B2A1AC670FF8227C5F8F17981F892" box="[421,447,1854,1873]" captionStart="Fig" captionStartId="4.[995,1025,149,166]" captionTargetBox="[102,968,150,977]" captionTargetId="figure-749@4.[100,970,148,979]" captionTargetPageId="4" captionText="Fig. 4. Light micrographs of female holotype MNHN- 655Ma (A, C-D), male paratype MNHN-661Ma (B, E), female paratypes MNHN-656Ma (H) and MNHN-657Ma (F), and additional specimens (G, I) of Echinoderes angelae sp. nov. showing trunk overview, details of mouth cone and neck, epibionts and ingested diatoms. A: Ventral trunk overview; B: mouth cone, showing the outer oral styles; C: dorsal neck view; D: ventral neck view; E: middorsal to laterodorsal view on right side of tergal plate of segments 10–11; F: lateral view of segment 11; G: ingested diatom; H: unknown type of epibionts attached to the posterior edge of segment 11; I: filamentous bacteria attached to the posterior edge of segment 11. Abbreviations: ba, bacteria; bs, basal sheath; dpl, dorsal placid; ep, end-piece; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; mvpl, midventral placid; ps, penile spines; te, tergal extension; tsp, trichoscalid plate." figureDoi="http://doi.org/10.5281/zenodo.10375301" httpUri="https://zenodo.org/record/10375301/files/figure.png" pageId="5" pageNumber="184">4F</figureCitation>
;
<figureCitation id="BE9B2A1AC670FF8227A6F8F179DBF892" box="[454,485,1854,1873]" captionStart="Fig" captionStartId="6.[997,1027,149,166]" captionTargetBox="[102,970,150,1260]" captionTargetId="figure-385@6.[100,972,148,1261]" captionTargetPageId="6" captionText="Fig. 6. Scanning electron micrographs of additional specimens of Echinoderes angelae sp. nov. showing trunk overview and details of introvert, neck, lateral terminal spines and sexually dimorphic features. A: Lateral trunk overview; B: introvert, sector 3; C: introvert, sector 4; D: ventral neck view; E: lateral terminal spine; F: male penile spines; G: female lateral terminal accessory spine. Abbreviations: ba, bacteria; ftz, fringed transverse zone; ltas, lateral terminal accessory spine; lts, lateral terminal spine; mvpl, midventral placid; ne, neck; ps, penile spines; psp, primary spinoscalid; sp, spinoscalid (followed by number of corresponding ring); te, tergal extension; ts, trichoscalid; tsp, trichoscalid plate; lambda marks (̂) indicate attachment point of spinoscalids." figureDoi="http://doi.org/10.5281/zenodo.10375307" httpUri="https://zenodo.org/record/10375307/files/figure.png" pageId="5" pageNumber="184">6G</figureCitation>
). Droplet-shaped sensory spots in subdorsal and ventromedial positions (
<figureCitation id="BE9B2A1AC670FF8222DCFA207D22F9C0" box="[1212,1308,1519,1539]" captionStart="Fig" captionStartId="3.[994,1024,149,166]" captionTargetBox="[101,967,149,1199]" captionTargetId="figure-599@3.[100,969,148,1201]" captionTargetPageId="3" captionText="Fig. 3. Line art drawing of Echinoderes angelae sp. nov. A: Ventral female overview; B: dorsal female overview; C: ventral male overview of segments 10–11; D: dorsal male overview of segments 10–11. Abbreviations: dpl, dorsal placid; go, gonopore; lat, lateral accessory tube; lddss, laterodorsal droplet-shaped sensory spot; ldmgco2, laterodorsal modified type 2 glandular cell outlet; ldrss, laterodorsal rounded sensory spot; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; lvgco1, lateroventral type 1 glandular cell outlet; lvmgco2, lateroventral modified type 2 glandular cell outlet; lvt, lateroventral tube; mddss, middorsal droplet-shaped sensory spot; mdgco1, middorsal type 1 glandular cell outlet; mds, middorsal spine; mldss, midlateral droplet-shaped sensory spot; mlmgco2, midlateral modified type 2 glandular cell outlet; ms, muscular scar; mvpl, midventral placid; ppf, primary pectinate fringe; ps, penile spines; S, segment (followed by number of corresponding segment); sddss, subdorsal droplet-shaped sensory spot; sdgco1, subdorsal type 1 glandular cell outlet; sdmgco2, subdorsal modified type 2 glandular cell outlet; sdrss, subdorsal rounded sensory spot; slne, sublateral nephridiopore; sls, sublateral spine; te, tergal extension; tsp, trichoscalid plate; vldss, ventrolateral droplet-shaped sensory spot; vmdss, ventromedial droplet-shaped sensory spot; vmgco1, ventromedial type 1 glandular cell outlet; vmrss, ventromedial rounded sensory spot." figureDoi="http://doi.org/10.5281/zenodo.10375299" httpUri="https://zenodo.org/record/10375299/files/figure.png" pageId="5" pageNumber="184">Fig. 3A–D</figureCitation>
;
<figureCitation id="BE9B2A1AC670FF822347FA207D7CF9C1" box="[1319,1346,1519,1538]" captionStart="Fig" captionStartId="7.[995,1025,149,166]" captionTargetBox="[102,968,150,1247]" captionTargetId="figure-378@7.[100,970,148,1249]" captionTargetPageId="7" captionText="Fig. 7. Scanning electron micrographs of additional specimens of Echinoderes angelae sp. nov. showing trunk segments and details of cuticular appendages. A: Subdorsal and laterodorsal sensory spots of segment 1, with the enlarged external ring of micropapillar (inset shows the intraspecific variability of these structures without such kind of external ring); B: ventral view of segments 2–6; C: subdorsal (top) and laterodorsal (bottom) modified type 2 glandular cell outlets of segments 2 and 4 respectively; D: laterodorsal view on left side of tergal plate of segment 2; E: nephridiopore and midlateral dropletshaped sensory spot of segment 9; F: subdorsal to midlateral view on right side of tergal plates of segments 8–9; G: middorsal spine of segment 4; H: subdorsal to midlateral view on left side of tergal plates of segments 3–4; I: laterodorsal tube of segment 10; J: lateral accessory tube of segment 8 (left) and sublateral spine of segment 7 (right); K: subdorsal type 1 glandular cell outlet of segment 9; L: ventral view of segments 10–11; M: subdorsal droplet-shaped sensory spot of segment 9. Abbreviations: go, gonopore; ms, muscular scar; spf, secondary pectinate fringe; modified type 2 glandular cell outlets are marked with continuous circles, and sensory spots with dashed teardrop-like figures." figureDoi="http://doi.org/10.5281/zenodo.10375309" httpUri="https://zenodo.org/record/10375309/files/figure.png" pageId="5" pageNumber="184">7L</figureCitation>
). Tergal extensions triangular, distally pointed, covered by hairs, distally elongated as hair-like extensions (
<figureCitation id="BE9B2A1AC670FF822597F9E87C69F9F9" box="[1015,1111,1575,1594]" captionStart="Fig" captionStartId="3.[994,1024,149,166]" captionTargetBox="[101,967,149,1199]" captionTargetId="figure-599@3.[100,969,148,1201]" captionTargetPageId="3" captionText="Fig. 3. Line art drawing of Echinoderes angelae sp. nov. A: Ventral female overview; B: dorsal female overview; C: ventral male overview of segments 10–11; D: dorsal male overview of segments 10–11. Abbreviations: dpl, dorsal placid; go, gonopore; lat, lateral accessory tube; lddss, laterodorsal droplet-shaped sensory spot; ldmgco2, laterodorsal modified type 2 glandular cell outlet; ldrss, laterodorsal rounded sensory spot; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; lvgco1, lateroventral type 1 glandular cell outlet; lvmgco2, lateroventral modified type 2 glandular cell outlet; lvt, lateroventral tube; mddss, middorsal droplet-shaped sensory spot; mdgco1, middorsal type 1 glandular cell outlet; mds, middorsal spine; mldss, midlateral droplet-shaped sensory spot; mlmgco2, midlateral modified type 2 glandular cell outlet; ms, muscular scar; mvpl, midventral placid; ppf, primary pectinate fringe; ps, penile spines; S, segment (followed by number of corresponding segment); sddss, subdorsal droplet-shaped sensory spot; sdgco1, subdorsal type 1 glandular cell outlet; sdmgco2, subdorsal modified type 2 glandular cell outlet; sdrss, subdorsal rounded sensory spot; slne, sublateral nephridiopore; sls, sublateral spine; te, tergal extension; tsp, trichoscalid plate; vldss, ventrolateral droplet-shaped sensory spot; vmdss, ventromedial droplet-shaped sensory spot; vmgco1, ventromedial type 1 glandular cell outlet; vmrss, ventromedial rounded sensory spot." figureDoi="http://doi.org/10.5281/zenodo.10375299" httpUri="https://zenodo.org/record/10375299/files/figure.png" pageId="5" pageNumber="184">Fig. 3A–D</figureCitation>
;
<figureCitation id="BE9B2A1AC670FF822201F9E87C45F9F9" box="[1121,1147,1575,1594]" captionStart="Fig" captionStartId="4.[995,1025,149,166]" captionTargetBox="[102,968,150,977]" captionTargetId="figure-749@4.[100,970,148,979]" captionTargetPageId="4" captionText="Fig. 4. Light micrographs of female holotype MNHN- 655Ma (A, C-D), male paratype MNHN-661Ma (B, E), female paratypes MNHN-656Ma (H) and MNHN-657Ma (F), and additional specimens (G, I) of Echinoderes angelae sp. nov. showing trunk overview, details of mouth cone and neck, epibionts and ingested diatoms. A: Ventral trunk overview; B: mouth cone, showing the outer oral styles; C: dorsal neck view; D: ventral neck view; E: middorsal to laterodorsal view on right side of tergal plate of segments 10–11; F: lateral view of segment 11; G: ingested diatom; H: unknown type of epibionts attached to the posterior edge of segment 11; I: filamentous bacteria attached to the posterior edge of segment 11. Abbreviations: ba, bacteria; bs, basal sheath; dpl, dorsal placid; ep, end-piece; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; mvpl, midventral placid; ps, penile spines; te, tergal extension; tsp, trichoscalid plate." figureDoi="http://doi.org/10.5281/zenodo.10375301" httpUri="https://zenodo.org/record/10375301/files/figure.png" pageId="5" pageNumber="184">4F</figureCitation>
;
<figureCitation id="BE9B2A1AC670FF8222E5F9E87C9FF9F9" box="[1157,1185,1575,1594]" captionStart="Fig" captionStartId="6.[997,1027,149,166]" captionTargetBox="[102,970,150,1260]" captionTargetId="figure-385@6.[100,972,148,1261]" captionTargetPageId="6" captionText="Fig. 6. Scanning electron micrographs of additional specimens of Echinoderes angelae sp. nov. showing trunk overview and details of introvert, neck, lateral terminal spines and sexually dimorphic features. A: Lateral trunk overview; B: introvert, sector 3; C: introvert, sector 4; D: ventral neck view; E: lateral terminal spine; F: male penile spines; G: female lateral terminal accessory spine. Abbreviations: ba, bacteria; ftz, fringed transverse zone; ltas, lateral terminal accessory spine; lts, lateral terminal spine; mvpl, midventral placid; ne, neck; ps, penile spines; psp, primary spinoscalid; sp, spinoscalid (followed by number of corresponding ring); te, tergal extension; ts, trichoscalid; tsp, trichoscalid plate; lambda marks (̂) indicate attachment point of spinoscalids." figureDoi="http://doi.org/10.5281/zenodo.10375307" httpUri="https://zenodo.org/record/10375307/files/figure.png" pageId="5" pageNumber="184">6E</figureCitation>
).
</paragraph>
<paragraph id="261F369FC670FF822552F9907CABF9B2" blockId="5.[818,1487,1630,1873]" box="[818,1173,1630,1650]" pageId="5" pageNumber="184">
<emphasis id="14D4EA8DC670FF822552F9907CABF9B2" bold="true" box="[818,1173,1630,1650]" italics="true" pageId="5" pageNumber="184">3.2.5. Notes on intraspecific variability</emphasis>
</paragraph>
<paragraph id="261F369FC670FF822531F9B57DF1F892" blockId="5.[818,1487,1630,1873]" pageId="5" pageNumber="184">
<taxonomicName id="E1A04D1CC670FF822531F9B57C36F94E" authority="Cepeda &amp; Gayet &amp; Spedicato &amp; Michaud &amp; Zeppilli, 2022" authorityName="Cepeda &amp; Gayet &amp; Spedicato &amp; Michaud &amp; Zeppilli" authorityYear="2022" box="[849,1032,1658,1677]" class="Cyclorhagida" family="Echinoderidae" genus="Echinoderes" kingdom="Animalia" order="Echinorhagata" pageId="5" pageNumber="184" phylum="Kinorhyncha" rank="species" species="angelae" status="sp. nov.">
<emphasis id="14D4EA8DC670FF822531F9B57C36F94E" bold="true" box="[849,1032,1658,1677]" italics="true" pageId="5" pageNumber="184">Echinoderes angelae</emphasis>
</taxonomicName>
<taxonomicNameLabel id="0FE757F6C670FF822271F9B47C5EF94D" box="[1041,1120,1659,1678]" pageId="5" pageNumber="184" rank="species">sp. nov.</taxonomicNameLabel>
shows certain degree of intraspecific variability not related to sex or sampling station. One of the most remarkable features of the species is the presence of three pairs of large sensory spots on segment 1 with a transversal row of conspicuously elongated hairs at the posterior edge of the papillae area (
<figureCitation id="BE9B2A1AC670FF82232CF9257DF5F93D" box="[1356,1483,1770,1790]" captionStart="Fig" captionStartId="3.[994,1024,149,166]" captionTargetBox="[101,967,149,1199]" captionTargetId="figure-599@3.[100,969,148,1201]" captionTargetPageId="3" captionText="Fig. 3. Line art drawing of Echinoderes angelae sp. nov. A: Ventral female overview; B: dorsal female overview; C: ventral male overview of segments 10–11; D: dorsal male overview of segments 10–11. Abbreviations: dpl, dorsal placid; go, gonopore; lat, lateral accessory tube; lddss, laterodorsal droplet-shaped sensory spot; ldmgco2, laterodorsal modified type 2 glandular cell outlet; ldrss, laterodorsal rounded sensory spot; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; lvgco1, lateroventral type 1 glandular cell outlet; lvmgco2, lateroventral modified type 2 glandular cell outlet; lvt, lateroventral tube; mddss, middorsal droplet-shaped sensory spot; mdgco1, middorsal type 1 glandular cell outlet; mds, middorsal spine; mldss, midlateral droplet-shaped sensory spot; mlmgco2, midlateral modified type 2 glandular cell outlet; ms, muscular scar; mvpl, midventral placid; ppf, primary pectinate fringe; ps, penile spines; S, segment (followed by number of corresponding segment); sddss, subdorsal droplet-shaped sensory spot; sdgco1, subdorsal type 1 glandular cell outlet; sdmgco2, subdorsal modified type 2 glandular cell outlet; sdrss, subdorsal rounded sensory spot; slne, sublateral nephridiopore; sls, sublateral spine; te, tergal extension; tsp, trichoscalid plate; vldss, ventrolateral droplet-shaped sensory spot; vmdss, ventromedial droplet-shaped sensory spot; vmgco1, ventromedial type 1 glandular cell outlet; vmrss, ventromedial rounded sensory spot." figureDoi="http://doi.org/10.5281/zenodo.10375299" httpUri="https://zenodo.org/record/10375299/files/figure.png" pageId="5" pageNumber="184">Fig. 3A and B</figureCitation>
;
<figureCitation id="BE9B2A1AC670FF822552F8C97B70F8DA" box="[818,846,1798,1817]" captionStart="Fig" captionStartId="5.[994,1024,149,166]" captionTargetBox="[102,967,150,1427]" captionTargetId="figure-287@5.[100,969,148,1429]" captionTargetPageId="5" captionText="Fig. 5. Light micrographs of female holotype MNHN- 655Ma (E), male paratype MNHN-661Ma (C–D), and additional specimens (A-B, F) of Echinoderes angelae sp. nov. showing trunk cuticular details. A: Middorsal to midlateral view on right side of tergal plate of segments 1–2; B: midlateral to ventromedial view on left side of cuticular plates of segments 5–7; C: ventral view of segments 5–6; D: dorsal view of segments 8–9; E: midlateral to ventromedial view on right side of cuticular plates of segments 9–10 (inset shows the intraspecific variability of the nephridiopore); F: midlateral to ventromedial view on right side of cuticular plates of segments 7–8. Abbreviations: lat, lateral accessory tube; lvt, lateroventral tube; ms, muscular scar; ppf, primary pectinate fringe; slne, sublateral nephridiopore; sls, sublateral spine; numbers after abbreviation indicate corresponding segment; glandular cell outlets are marked with continuous circles, and sensory spots with dashed circles or teardrop-like figures." figureDoi="http://doi.org/10.5281/zenodo.10375305" httpUri="https://zenodo.org/record/10375305/files/figure.png" pageId="5" pageNumber="184">5A</figureCitation>
;
<figureCitation id="BE9B2A1AC670FF822539F8C97B46F8DA" box="[857,888,1798,1817]" captionStart="Fig" captionStartId="7.[995,1025,149,166]" captionTargetBox="[102,968,150,1247]" captionTargetId="figure-378@7.[100,970,148,1249]" captionTargetPageId="7" captionText="Fig. 7. Scanning electron micrographs of additional specimens of Echinoderes angelae sp. nov. showing trunk segments and details of cuticular appendages. A: Subdorsal and laterodorsal sensory spots of segment 1, with the enlarged external ring of micropapillar (inset shows the intraspecific variability of these structures without such kind of external ring); B: ventral view of segments 2–6; C: subdorsal (top) and laterodorsal (bottom) modified type 2 glandular cell outlets of segments 2 and 4 respectively; D: laterodorsal view on left side of tergal plate of segment 2; E: nephridiopore and midlateral dropletshaped sensory spot of segment 9; F: subdorsal to midlateral view on right side of tergal plates of segments 8–9; G: middorsal spine of segment 4; H: subdorsal to midlateral view on left side of tergal plates of segments 3–4; I: laterodorsal tube of segment 10; J: lateral accessory tube of segment 8 (left) and sublateral spine of segment 7 (right); K: subdorsal type 1 glandular cell outlet of segment 9; L: ventral view of segments 10–11; M: subdorsal droplet-shaped sensory spot of segment 9. Abbreviations: go, gonopore; ms, muscular scar; spf, secondary pectinate fringe; modified type 2 glandular cell outlets are marked with continuous circles, and sensory spots with dashed teardrop-like figures." figureDoi="http://doi.org/10.5281/zenodo.10375309" httpUri="https://zenodo.org/record/10375309/files/figure.png" pageId="5" pageNumber="184">7A</figureCitation>
). However,
<specimenCount id="30A6FD16C670FF822589F8C87CBBF8DA" box="[1001,1157,1798,1818]" count="7" pageId="5" pageNumber="184" type="generic">seven specimens</specimenCount>
mounted for SEM (
<specimenCount id="30A6FD16C670FF822320F8C97D9BF8DA" box="[1344,1445,1798,1817]" count="4" pageId="5" pageNumber="184" type="male">four males</specimenCount>
and
<specimenCount id="30A6FD16C670FF822552F8ED7B8DF8F6" box="[818,947,1826,1845]" count="3" pageId="5" pageNumber="184" type="female">three females</specimenCount>
) showed these sensory spots without such kind of hairs (
<figureCitation id="BE9B2A1AC670FF82255AF8F17BBCF892" box="[826,898,1854,1873]" captionStart="Fig" captionStartId="7.[995,1025,149,166]" captionTargetBox="[102,968,150,1247]" captionTargetId="figure-378@7.[100,970,148,1249]" captionTargetPageId="7" captionText="Fig. 7. Scanning electron micrographs of additional specimens of Echinoderes angelae sp. nov. showing trunk segments and details of cuticular appendages. A: Subdorsal and laterodorsal sensory spots of segment 1, with the enlarged external ring of micropapillar (inset shows the intraspecific variability of these structures without such kind of external ring); B: ventral view of segments 2–6; C: subdorsal (top) and laterodorsal (bottom) modified type 2 glandular cell outlets of segments 2 and 4 respectively; D: laterodorsal view on left side of tergal plate of segment 2; E: nephridiopore and midlateral dropletshaped sensory spot of segment 9; F: subdorsal to midlateral view on right side of tergal plates of segments 8–9; G: middorsal spine of segment 4; H: subdorsal to midlateral view on left side of tergal plates of segments 3–4; I: laterodorsal tube of segment 10; J: lateral accessory tube of segment 8 (left) and sublateral spine of segment 7 (right); K: subdorsal type 1 glandular cell outlet of segment 9; L: ventral view of segments 10–11; M: subdorsal droplet-shaped sensory spot of segment 9. Abbreviations: go, gonopore; ms, muscular scar; spf, secondary pectinate fringe; modified type 2 glandular cell outlets are marked with continuous circles, and sensory spots with dashed teardrop-like figures." figureDoi="http://doi.org/10.5281/zenodo.10375309" httpUri="https://zenodo.org/record/10375309/files/figure.png" pageId="5" pageNumber="184">Fig. 7A</figureCitation>
). Neither juveniles of the species possess the elongated hairs,
</paragraph>
<caption id="72DF6617C673FF812585FF5A7D3FFE31" ID-DOI="http://doi.org/10.5281/zenodo.10375307" ID-Zenodo-Dep="10375307" httpUri="https://zenodo.org/record/10375307/files/figure.png" pageId="6" pageNumber="185" startId="6.[997,1027,149,166]" targetBox="[102,970,150,1260]" targetPageId="6" targetType="figure">
<paragraph id="261F369FC673FF812585FF5A7D3FFE31" blockId="6.[997,1488,149,498]" pageId="6" pageNumber="185">
<emphasis id="14D4EA8DC673FF812585FF5A7C19FF65" bold="true" box="[997,1063,149,166]" pageId="6" pageNumber="185">Fig. 6.</emphasis>
Scanning electron micrographs of additional specimens of
<taxonomicName id="E1A04D1CC673FF81223CFF607CC1FF03" authority="Cepeda &amp; Gayet &amp; Spedicato &amp; Michaud &amp; Zeppilli, 2022" authorityName="Cepeda &amp; Gayet &amp; Spedicato &amp; Michaud &amp; Zeppilli" authorityYear="2022" box="[1116,1279,175,192]" class="Cyclorhagida" family="Echinoderidae" genus="Echinoderes" kingdom="Animalia" order="Echinorhagata" pageId="6" pageNumber="185" phylum="Kinorhyncha" rank="species" species="angelae" status="sp. nov.">
<emphasis id="14D4EA8DC673FF81223CFF607CC1FF03" bold="true" box="[1116,1279,175,192]" italics="true" pageId="6" pageNumber="185">Echinoderes angelae</emphasis>
</taxonomicName>
<taxonomicNameLabel id="0FE757F6C673FF812367FF607D72FF7C" box="[1287,1356,175,191]" pageId="6" pageNumber="185" rank="species">sp. nov.</taxonomicNameLabel>
showing trunk overview and details of introvert, neck, lateral terminal spines and sexually dimorphic features. A: Lateral trunk overview; B: introvert, sector 3; C: introvert, sector 4; D: ventral neck view; E: lateral terminal spine; F: male penile spines; G: female lateral terminal accessory spine. Abbreviations: ba, bacteria; ftz, fringed transverse zone; ltas, lateral terminal accessory spine; lts, lateral terminal spine; mvpl, midventral placid; ne, neck; ps, penile spines; psp, primary spinoscalid; sp, spinoscalid (followed by number of corresponding ring); te, tergal extension; ts, trichoscalid; tsp, trichoscalid plate; lambda marks (̂) indicate attachment point of spinoscalids.
</paragraph>
</caption>
<paragraph id="261F369FC673FF812604FAA879BBFA54" blockId="6.[100,770,1383,1877]" pageId="6" pageNumber="185">which suggests that the observed specimens could be pre-adults that did not develop the structures yet.</paragraph>
<paragraph id="261F369FC673FF8126E4FA507AB1F99A" blockId="6.[100,770,1383,1877]" pageId="6" pageNumber="185">
Shape and position of nephridiopores also showed variation among the studied specimens. This structure varies from completely oval or slightly triangular, located near the anterior margin of segment 9 without rows of hairs on top (ca. 70% of the examined specimens) to more buttonhole-shaped, posteriorly displaced having a few rows of hairs on top (ca. 30% of the examined specimens) (
<figureCitation id="BE9B2A1AC673FF81242FF9E57AAAF9FD" box="[591,660,1578,1598]" captionStart="Fig" captionStartId="5.[994,1024,149,166]" captionTargetBox="[102,967,150,1427]" captionTargetId="figure-287@5.[100,969,148,1429]" captionTargetPageId="5" captionText="Fig. 5. Light micrographs of female holotype MNHN- 655Ma (E), male paratype MNHN-661Ma (C–D), and additional specimens (A-B, F) of Echinoderes angelae sp. nov. showing trunk cuticular details. A: Middorsal to midlateral view on right side of tergal plate of segments 1–2; B: midlateral to ventromedial view on left side of cuticular plates of segments 5–7; C: ventral view of segments 5–6; D: dorsal view of segments 8–9; E: midlateral to ventromedial view on right side of cuticular plates of segments 9–10 (inset shows the intraspecific variability of the nephridiopore); F: midlateral to ventromedial view on right side of cuticular plates of segments 7–8. Abbreviations: lat, lateral accessory tube; lvt, lateroventral tube; ms, muscular scar; ppf, primary pectinate fringe; slne, sublateral nephridiopore; sls, sublateral spine; numbers after abbreviation indicate corresponding segment; glandular cell outlets are marked with continuous circles, and sensory spots with dashed circles or teardrop-like figures." figureDoi="http://doi.org/10.5281/zenodo.10375305" httpUri="https://zenodo.org/record/10375305/files/figure.png" pageId="6" pageNumber="185">Fig. 5E</figureCitation>
;
<figureCitation id="BE9B2A1AC673FF8124FEF9E47A84F9FD" box="[670,698,1579,1598]" captionStart="Fig" captionStartId="7.[995,1025,149,166]" captionTargetBox="[102,968,150,1247]" captionTargetId="figure-378@7.[100,970,148,1249]" captionTargetPageId="7" captionText="Fig. 7. Scanning electron micrographs of additional specimens of Echinoderes angelae sp. nov. showing trunk segments and details of cuticular appendages. A: Subdorsal and laterodorsal sensory spots of segment 1, with the enlarged external ring of micropapillar (inset shows the intraspecific variability of these structures without such kind of external ring); B: ventral view of segments 2–6; C: subdorsal (top) and laterodorsal (bottom) modified type 2 glandular cell outlets of segments 2 and 4 respectively; D: laterodorsal view on left side of tergal plate of segment 2; E: nephridiopore and midlateral dropletshaped sensory spot of segment 9; F: subdorsal to midlateral view on right side of tergal plates of segments 8–9; G: middorsal spine of segment 4; H: subdorsal to midlateral view on left side of tergal plates of segments 3–4; I: laterodorsal tube of segment 10; J: lateral accessory tube of segment 8 (left) and sublateral spine of segment 7 (right); K: subdorsal type 1 glandular cell outlet of segment 9; L: ventral view of segments 10–11; M: subdorsal droplet-shaped sensory spot of segment 9. Abbreviations: go, gonopore; ms, muscular scar; spf, secondary pectinate fringe; modified type 2 glandular cell outlets are marked with continuous circles, and sensory spots with dashed teardrop-like figures." figureDoi="http://doi.org/10.5281/zenodo.10375309" httpUri="https://zenodo.org/record/10375309/files/figure.png" pageId="6" pageNumber="185">7E</figureCitation>
). These differences were observed in both LM and SEM specimens.
</paragraph>
<paragraph id="261F369FC673FF8126E4F9AD7C76F9E2" blockId="6.[100,770,1383,1877]" lastBlockId="6.[818,1488,1383,1570]" pageId="6" pageNumber="185">
Segments 10 and 11 also showed conspicuous intraspecific variability regarding the length of the tergal extensions and the primary pectinate fringe. Thus, some specimens had shorter tergal extensions, without the hair-like distal end, as well as a primary pectinate fringe with barely elongated tips (ca. 20% of the examined specimens) (
<figureCitation id="BE9B2A1AC673FF81260CF921788EF8C2" box="[108,176,1774,1793]" captionStart="Fig" captionStartId="6.[997,1027,149,166]" captionTargetBox="[102,970,150,1260]" captionTargetId="figure-385@6.[100,972,148,1261]" captionTargetPageId="6" captionText="Fig. 6. Scanning electron micrographs of additional specimens of Echinoderes angelae sp. nov. showing trunk overview and details of introvert, neck, lateral terminal spines and sexually dimorphic features. A: Lateral trunk overview; B: introvert, sector 3; C: introvert, sector 4; D: ventral neck view; E: lateral terminal spine; F: male penile spines; G: female lateral terminal accessory spine. Abbreviations: ba, bacteria; ftz, fringed transverse zone; ltas, lateral terminal accessory spine; lts, lateral terminal spine; mvpl, midventral placid; ne, neck; ps, penile spines; psp, primary spinoscalid; sp, spinoscalid (followed by number of corresponding ring); te, tergal extension; ts, trichoscalid; tsp, trichoscalid plate; lambda marks (̂) indicate attachment point of spinoscalids." figureDoi="http://doi.org/10.5281/zenodo.10375307" httpUri="https://zenodo.org/record/10375307/files/figure.png" pageId="6" pageNumber="185">Fig. 6E</figureCitation>
). Others had tergal extensions with an elongated, hair-like distal end and primary pectinate fringes with longer tips (ca. 80% of the examined specimens) (
<figureCitation id="BE9B2A1AC673FF812721F8EA799CF8FA" box="[321,418,1829,1849]" captionStart="Fig" captionStartId="3.[994,1024,149,166]" captionTargetBox="[101,967,149,1199]" captionTargetId="figure-599@3.[100,969,148,1201]" captionTargetPageId="3" captionText="Fig. 3. Line art drawing of Echinoderes angelae sp. nov. A: Ventral female overview; B: dorsal female overview; C: ventral male overview of segments 10–11; D: dorsal male overview of segments 10–11. Abbreviations: dpl, dorsal placid; go, gonopore; lat, lateral accessory tube; lddss, laterodorsal droplet-shaped sensory spot; ldmgco2, laterodorsal modified type 2 glandular cell outlet; ldrss, laterodorsal rounded sensory spot; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; lvgco1, lateroventral type 1 glandular cell outlet; lvmgco2, lateroventral modified type 2 glandular cell outlet; lvt, lateroventral tube; mddss, middorsal droplet-shaped sensory spot; mdgco1, middorsal type 1 glandular cell outlet; mds, middorsal spine; mldss, midlateral droplet-shaped sensory spot; mlmgco2, midlateral modified type 2 glandular cell outlet; ms, muscular scar; mvpl, midventral placid; ppf, primary pectinate fringe; ps, penile spines; S, segment (followed by number of corresponding segment); sddss, subdorsal droplet-shaped sensory spot; sdgco1, subdorsal type 1 glandular cell outlet; sdmgco2, subdorsal modified type 2 glandular cell outlet; sdrss, subdorsal rounded sensory spot; slne, sublateral nephridiopore; sls, sublateral spine; te, tergal extension; tsp, trichoscalid plate; vldss, ventrolateral droplet-shaped sensory spot; vmdss, ventromedial droplet-shaped sensory spot; vmgco1, ventromedial type 1 glandular cell outlet; vmrss, ventromedial rounded sensory spot." figureDoi="http://doi.org/10.5281/zenodo.10375299" httpUri="https://zenodo.org/record/10375299/files/figure.png" pageId="6" pageNumber="185">Fig. 3A–D</figureCitation>
;
<figureCitation id="BE9B2A1AC673FF8127CFF8E97984F8FA" box="[431,442,1830,1849]" captionStart="Fig" captionStartId="4.[995,1025,149,166]" captionTargetBox="[102,968,150,977]" captionTargetId="figure-749@4.[100,970,148,979]" captionTargetPageId="4" captionText="Fig. 4. Light micrographs of female holotype MNHN- 655Ma (A, C-D), male paratype MNHN-661Ma (B, E), female paratypes MNHN-656Ma (H) and MNHN-657Ma (F), and additional specimens (G, I) of Echinoderes angelae sp. nov. showing trunk overview, details of mouth cone and neck, epibionts and ingested diatoms. A: Ventral trunk overview; B: mouth cone, showing the outer oral styles; C: dorsal neck view; D: ventral neck view; E: middorsal to laterodorsal view on right side of tergal plate of segments 10–11; F: lateral view of segment 11; G: ingested diatom; H: unknown type of epibionts attached to the posterior edge of segment 11; I: filamentous bacteria attached to the posterior edge of segment 11. Abbreviations: ba, bacteria; bs, basal sheath; dpl, dorsal placid; ep, end-piece; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; mvpl, midventral placid; ps, penile spines; te, tergal extension; tsp, trichoscalid plate." figureDoi="http://doi.org/10.5281/zenodo.10375301" httpUri="https://zenodo.org/record/10375301/files/figure.png" pageId="6" pageNumber="185">4</figureCitation>
E-F; 6F-G). The primary pectinate fringe of segment 10 also showed some intraspecific variation concerning the length of the tips at the ventrolateral region, with ca. 30% of the examined specimens showing conspicuously elongated tips (reaching the edge of segment 11) at this area. Finally, the relative length of the lateral terminal spines also showed intraspecific variability (LTS:TL ratio of 6–10% in 15 out of 52 measured specimens vs. 15–25% in 37 out of 52 measured specimens). These differences were observed in both LM and SEM specimens.
</paragraph>
<paragraph id="261F369FC673FF812552F9897BCAF99A" blockId="6.[818,1487,1606,1877]" box="[818,1012,1606,1625]" pageId="6" pageNumber="185">
<emphasis id="14D4EA8DC673FF812552F9897BCAF99A" bold="true" box="[818,1012,1606,1625]" italics="true" pageId="6" pageNumber="185">3.2.6. Other remarks</emphasis>
</paragraph>
<paragraph id="261F369FC673FF802531F9AD793CFA97" blockId="6.[818,1487,1606,1877]" lastBlockId="7.[100,771,1345,1504]" lastPageId="7" lastPageNumber="186" pageId="6" pageNumber="185">
Several specimens (ca. 15% of the total number) had epibionts. The most common ones were filamentous bacteria of different lengths (
<figureCitation id="BE9B2A1AC673FF81255AF9557B45F96E" box="[826,891,1690,1709]" captionStart="Fig" captionStartId="4.[995,1025,149,166]" captionTargetBox="[102,968,150,977]" captionTargetId="figure-749@4.[100,970,148,979]" captionTargetPageId="4" captionText="Fig. 4. Light micrographs of female holotype MNHN- 655Ma (A, C-D), male paratype MNHN-661Ma (B, E), female paratypes MNHN-656Ma (H) and MNHN-657Ma (F), and additional specimens (G, I) of Echinoderes angelae sp. nov. showing trunk overview, details of mouth cone and neck, epibionts and ingested diatoms. A: Ventral trunk overview; B: mouth cone, showing the outer oral styles; C: dorsal neck view; D: ventral neck view; E: middorsal to laterodorsal view on right side of tergal plate of segments 10–11; F: lateral view of segment 11; G: ingested diatom; H: unknown type of epibionts attached to the posterior edge of segment 11; I: filamentous bacteria attached to the posterior edge of segment 11. Abbreviations: ba, bacteria; bs, basal sheath; dpl, dorsal placid; ep, end-piece; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; mvpl, midventral placid; ps, penile spines; te, tergal extension; tsp, trichoscalid plate." figureDoi="http://doi.org/10.5281/zenodo.10375301" httpUri="https://zenodo.org/record/10375301/files/figure.png" pageId="6" pageNumber="185">Fig. 4I</figureCitation>
;
<figureCitation id="BE9B2A1AC673FF8125E9F9557B98F96E" box="[905,934,1690,1709]" captionStart="Fig" captionStartId="6.[997,1027,149,166]" captionTargetBox="[102,970,150,1260]" captionTargetId="figure-385@6.[100,972,148,1261]" captionTargetPageId="6" captionText="Fig. 6. Scanning electron micrographs of additional specimens of Echinoderes angelae sp. nov. showing trunk overview and details of introvert, neck, lateral terminal spines and sexually dimorphic features. A: Lateral trunk overview; B: introvert, sector 3; C: introvert, sector 4; D: ventral neck view; E: lateral terminal spine; F: male penile spines; G: female lateral terminal accessory spine. Abbreviations: ba, bacteria; ftz, fringed transverse zone; ltas, lateral terminal accessory spine; lts, lateral terminal spine; mvpl, midventral placid; ne, neck; ps, penile spines; psp, primary spinoscalid; sp, spinoscalid (followed by number of corresponding ring); te, tergal extension; ts, trichoscalid; tsp, trichoscalid plate; lambda marks (̂) indicate attachment point of spinoscalids." figureDoi="http://doi.org/10.5281/zenodo.10375307" httpUri="https://zenodo.org/record/10375307/files/figure.png" pageId="6" pageNumber="185">6G</figureCitation>
;
<figureCitation id="BE9B2A1AC673FF8125D3F9557B82F96E" box="[947,956,1690,1709]" captionStart="Fig" captionStartId="8.[997,1027,149,166]" captionTargetBox="[102,970,150,1291]" captionTargetId="figure-503@8.[100,972,148,1293]" captionTargetPageId="8" captionText="Fig. 8. Scanning electron micrographs showing two types of prokaryotic (likely bacteria) epibionts found on additional specimens of Echinoderes angelae sp. nov. A: Sternal plates of segments 6–7 with filamentous bacteria; B: sternal plates of segments 6–8 with filamentous bacteria; C: filamentous bacteria attached to the ventromedial droplet-shaped sensory spot of segment 6; D: bacteria near the ventrolateral droplet-shaped sensory spot of segment 2; E: bacteria on the muscular scar of segment 6; F: filamentous bacteria attached to the subdorsal enlarged sensory spot of segment 1; G: sternal plates of segments 9–11 and lateral terminal spines with filamentous bacteria; H: bacteria near the lateroventral tube of segment 5; I: bacteria and filamentous bacteria on nephridiopore of segment 9. Abbreviations: tu, tube; arrows indicate the presence and/or attachment point of bacteria and filamentous bacteria respectively." figureDoi="http://doi.org/10.5281/zenodo.10375311" httpUri="https://zenodo.org/record/10375311/files/figure.png" pageId="6" pageNumber="185">8</figureCitation>
A-C, F-G), but unicellular bacteria were also observed (
<figureCitation id="BE9B2A1AC673FF81255AF9797BA6F90A" box="[826,920,1718,1737]" captionStart="Fig" captionStartId="8.[997,1027,149,166]" captionTargetBox="[102,970,150,1291]" captionTargetId="figure-503@8.[100,972,148,1293]" captionTargetPageId="8" captionText="Fig. 8. Scanning electron micrographs showing two types of prokaryotic (likely bacteria) epibionts found on additional specimens of Echinoderes angelae sp. nov. A: Sternal plates of segments 6–7 with filamentous bacteria; B: sternal plates of segments 6–8 with filamentous bacteria; C: filamentous bacteria attached to the ventromedial droplet-shaped sensory spot of segment 6; D: bacteria near the ventrolateral droplet-shaped sensory spot of segment 2; E: bacteria on the muscular scar of segment 6; F: filamentous bacteria attached to the subdorsal enlarged sensory spot of segment 1; G: sternal plates of segments 9–11 and lateral terminal spines with filamentous bacteria; H: bacteria near the lateroventral tube of segment 5; I: bacteria and filamentous bacteria on nephridiopore of segment 9. Abbreviations: tu, tube; arrows indicate the presence and/or attachment point of bacteria and filamentous bacteria respectively." figureDoi="http://doi.org/10.5281/zenodo.10375311" httpUri="https://zenodo.org/record/10375311/files/figure.png" pageId="6" pageNumber="185">Fig. 8D–E</figureCitation>
, H-I). Finally, unknown, rod-shaped epibionts, with a distal constriction, were also detected (
<figureCitation id="BE9B2A1AC673FF8122EEF91D7CDEF926" box="[1166,1248,1746,1765]" captionStart="Fig" captionStartId="4.[995,1025,149,166]" captionTargetBox="[102,968,150,977]" captionTargetId="figure-749@4.[100,970,148,979]" captionTargetPageId="4" captionText="Fig. 4. Light micrographs of female holotype MNHN- 655Ma (A, C-D), male paratype MNHN-661Ma (B, E), female paratypes MNHN-656Ma (H) and MNHN-657Ma (F), and additional specimens (G, I) of Echinoderes angelae sp. nov. showing trunk overview, details of mouth cone and neck, epibionts and ingested diatoms. A: Ventral trunk overview; B: mouth cone, showing the outer oral styles; C: dorsal neck view; D: ventral neck view; E: middorsal to laterodorsal view on right side of tergal plate of segments 10–11; F: lateral view of segment 11; G: ingested diatom; H: unknown type of epibionts attached to the posterior edge of segment 11; I: filamentous bacteria attached to the posterior edge of segment 11. Abbreviations: ba, bacteria; bs, basal sheath; dpl, dorsal placid; ep, end-piece; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; mvpl, midventral placid; ps, penile spines; te, tergal extension; tsp, trichoscalid plate." figureDoi="http://doi.org/10.5281/zenodo.10375301" httpUri="https://zenodo.org/record/10375301/files/figure.png" pageId="6" pageNumber="185">Fig. 4H</figureCitation>
). Epibionts were more commonly found attached to the ventral surface of the posterior segments, the nephridiopore and the lateral terminal spines (
<figureCitation id="BE9B2A1AC673FF81232FF8C57DF5F8DE" box="[1359,1483,1802,1821]" captionStart="Fig" captionStartId="4.[995,1025,149,166]" captionTargetBox="[102,968,150,977]" captionTargetId="figure-749@4.[100,970,148,979]" captionTargetPageId="4" captionText="Fig. 4. Light micrographs of female holotype MNHN- 655Ma (A, C-D), male paratype MNHN-661Ma (B, E), female paratypes MNHN-656Ma (H) and MNHN-657Ma (F), and additional specimens (G, I) of Echinoderes angelae sp. nov. showing trunk overview, details of mouth cone and neck, epibionts and ingested diatoms. A: Ventral trunk overview; B: mouth cone, showing the outer oral styles; C: dorsal neck view; D: ventral neck view; E: middorsal to laterodorsal view on right side of tergal plate of segments 10–11; F: lateral view of segment 11; G: ingested diatom; H: unknown type of epibionts attached to the posterior edge of segment 11; I: filamentous bacteria attached to the posterior edge of segment 11. Abbreviations: ba, bacteria; bs, basal sheath; dpl, dorsal placid; ep, end-piece; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; mvpl, midventral placid; ps, penile spines; te, tergal extension; tsp, trichoscalid plate." figureDoi="http://doi.org/10.5281/zenodo.10375301" httpUri="https://zenodo.org/record/10375301/files/figure.png" pageId="6" pageNumber="185">Fig. 4H and I</figureCitation>
;
<figureCitation id="BE9B2A1AC673FF812552F8EA7B71F8FB" box="[818,847,1829,1848]" captionStart="Fig" captionStartId="6.[997,1027,149,166]" captionTargetBox="[102,970,150,1260]" captionTargetId="figure-385@6.[100,972,148,1261]" captionTargetPageId="6" captionText="Fig. 6. Scanning electron micrographs of additional specimens of Echinoderes angelae sp. nov. showing trunk overview and details of introvert, neck, lateral terminal spines and sexually dimorphic features. A: Lateral trunk overview; B: introvert, sector 3; C: introvert, sector 4; D: ventral neck view; E: lateral terminal spine; F: male penile spines; G: female lateral terminal accessory spine. Abbreviations: ba, bacteria; ftz, fringed transverse zone; ltas, lateral terminal accessory spine; lts, lateral terminal spine; mvpl, midventral placid; ne, neck; ps, penile spines; psp, primary spinoscalid; sp, spinoscalid (followed by number of corresponding ring); te, tergal extension; ts, trichoscalid; tsp, trichoscalid plate; lambda marks (̂) indicate attachment point of spinoscalids." figureDoi="http://doi.org/10.5281/zenodo.10375307" httpUri="https://zenodo.org/record/10375307/files/figure.png" pageId="6" pageNumber="185">6G</figureCitation>
;
<figureCitation id="BE9B2A1AC673FF81253AF8E97B49F8FA" box="[858,887,1830,1849]" captionStart="Fig" captionStartId="8.[997,1027,149,166]" captionTargetBox="[102,970,150,1291]" captionTargetId="figure-503@8.[100,972,148,1293]" captionTargetPageId="8" captionText="Fig. 8. Scanning electron micrographs showing two types of prokaryotic (likely bacteria) epibionts found on additional specimens of Echinoderes angelae sp. nov. A: Sternal plates of segments 6–7 with filamentous bacteria; B: sternal plates of segments 6–8 with filamentous bacteria; C: filamentous bacteria attached to the ventromedial droplet-shaped sensory spot of segment 6; D: bacteria near the ventrolateral droplet-shaped sensory spot of segment 2; E: bacteria on the muscular scar of segment 6; F: filamentous bacteria attached to the subdorsal enlarged sensory spot of segment 1; G: sternal plates of segments 9–11 and lateral terminal spines with filamentous bacteria; H: bacteria near the lateroventral tube of segment 5; I: bacteria and filamentous bacteria on nephridiopore of segment 9. Abbreviations: tu, tube; arrows indicate the presence and/or attachment point of bacteria and filamentous bacteria respectively." figureDoi="http://doi.org/10.5281/zenodo.10375311" httpUri="https://zenodo.org/record/10375311/files/figure.png" pageId="6" pageNumber="185">8G</figureCitation>
), but some of them were also present at the anterior segments near sensory spots, tubes, muscular scars and glandular cell outlets (
<figureCitation id="BE9B2A1AC672FF80260CFA8E78C8FA97" box="[108,246,1345,1365]" captionStart="Fig" captionStartId="8.[997,1027,149,166]" captionTargetBox="[102,970,150,1291]" captionTargetId="figure-503@8.[100,972,148,1293]" captionTargetPageId="8" captionText="Fig. 8. Scanning electron micrographs showing two types of prokaryotic (likely bacteria) epibionts found on additional specimens of Echinoderes angelae sp. nov. A: Sternal plates of segments 6–7 with filamentous bacteria; B: sternal plates of segments 6–8 with filamentous bacteria; C: filamentous bacteria attached to the ventromedial droplet-shaped sensory spot of segment 6; D: bacteria near the ventrolateral droplet-shaped sensory spot of segment 2; E: bacteria on the muscular scar of segment 6; F: filamentous bacteria attached to the subdorsal enlarged sensory spot of segment 1; G: sternal plates of segments 9–11 and lateral terminal spines with filamentous bacteria; H: bacteria near the lateroventral tube of segment 5; I: bacteria and filamentous bacteria on nephridiopore of segment 9. Abbreviations: tu, tube; arrows indicate the presence and/or attachment point of bacteria and filamentous bacteria respectively." figureDoi="http://doi.org/10.5281/zenodo.10375311" httpUri="https://zenodo.org/record/10375311/files/figure.png" pageId="7" pageNumber="186">Fig. 8A–F, H–I</figureCitation>
).
</paragraph>
<caption id="72DF6617C672FF802583FF5A7CE7FD32" ID-DOI="http://doi.org/10.5281/zenodo.10375309" ID-Zenodo-Dep="10375309" httpUri="https://zenodo.org/record/10375309/files/figure.png" pageId="7" pageNumber="186" startId="7.[995,1025,149,166]" targetBox="[102,968,150,1247]" targetPageId="7" targetType="figure">
<paragraph id="261F369FC672FF802583FF5A7CE7FD32" blockId="7.[995,1488,149,753]" pageId="7" pageNumber="186">
<emphasis id="14D4EA8DC672FF802583FF5A7C18FF65" bold="true" box="[995,1062,149,166]" pageId="7" pageNumber="186">Fig. 7.</emphasis>
Scanning electron micrographs of additional specimens of
<taxonomicName id="E1A04D1CC672FF80223AFF607CC3FF03" authority="Cepeda &amp; Gayet &amp; Spedicato &amp; Michaud &amp; Zeppilli, 2022" authorityName="Cepeda &amp; Gayet &amp; Spedicato &amp; Michaud &amp; Zeppilli" authorityYear="2022" box="[1114,1277,175,192]" class="Cyclorhagida" family="Echinoderidae" genus="Echinoderes" kingdom="Animalia" order="Echinorhagata" pageId="7" pageNumber="186" phylum="Kinorhyncha" rank="species" species="angelae" status="sp. nov.">
<emphasis id="14D4EA8DC672FF80223AFF607CC3FF03" bold="true" box="[1114,1277,175,192]" italics="true" pageId="7" pageNumber="186">Echinoderes angelae</emphasis>
</taxonomicName>
<taxonomicNameLabel id="0FE757F6C672FF802365FF607D75FF7C" box="[1285,1355,175,191]" pageId="7" pageNumber="186" rank="species">sp. nov.</taxonomicNameLabel>
showing trunk segments and details of cuticular appendages. A: Subdorsal and laterodorsal sensory spots of segment 1, with the enlarged external ring of micropapillar (inset shows the intraspecific variability of these structures without such kind of external ring); B: ventral view of segments 2–6; C: subdorsal (top) and laterodorsal (bottom) modified type 2 glandular cell outlets of segments 2 and 4 respectively; D: laterodorsal view on left side of tergal plate of segment 2; E: nephridiopore and midlateral dropletshaped sensory spot of segment 9; F: subdorsal to midlateral view on right side of tergal plates of segments 8–9; G: middorsal spine of segment 4; H: subdorsal to midlateral view on left side of tergal plates of segments 3–4; I: laterodorsal tube of segment 10; J: lateral accessory tube of segment 8 (left) and sublateral spine of segment 7 (right); K: subdorsal type 1 glandular cell outlet of segment 9; L: ventral view of segments 10–11; M: subdorsal droplet-shaped sensory spot of segment 9. Abbreviations: go, gonopore; ms, muscular scar; spf, secondary pectinate fringe; modified type 2 glandular cell outlets are marked with continuous circles, and sensory spots with dashed teardrop-like figures.
</paragraph>
</caption>
<paragraph id="261F369FC672FF8026E4FA927902FA23" blockId="7.[100,771,1345,1504]" pageId="7" pageNumber="186">
Some specimens (ca. 10% of the total number) had ingested clusters of diatoms at the gut, and even though their frustules seemed to be unaltered, a partially digested, green content was observed (
<figureCitation id="BE9B2A1AC672FF8024CEFA5A7AC6FA6B" box="[686,760,1429,1448]" captionStart="Fig" captionStartId="4.[995,1025,149,166]" captionTargetBox="[102,968,150,977]" captionTargetId="figure-749@4.[100,970,148,979]" captionTargetPageId="4" captionText="Fig. 4. Light micrographs of female holotype MNHN- 655Ma (A, C-D), male paratype MNHN-661Ma (B, E), female paratypes MNHN-656Ma (H) and MNHN-657Ma (F), and additional specimens (G, I) of Echinoderes angelae sp. nov. showing trunk overview, details of mouth cone and neck, epibionts and ingested diatoms. A: Ventral trunk overview; B: mouth cone, showing the outer oral styles; C: dorsal neck view; D: ventral neck view; E: middorsal to laterodorsal view on right side of tergal plate of segments 10–11; F: lateral view of segment 11; G: ingested diatom; H: unknown type of epibionts attached to the posterior edge of segment 11; I: filamentous bacteria attached to the posterior edge of segment 11. Abbreviations: ba, bacteria; bs, basal sheath; dpl, dorsal placid; ep, end-piece; ldt, laterodorsal tube; ltas, lateral terminal accessory spine; lts, lateral terminal spine; mvpl, midventral placid; ps, penile spines; te, tergal extension; tsp, trichoscalid plate." figureDoi="http://doi.org/10.5281/zenodo.10375301" httpUri="https://zenodo.org/record/10375301/files/figure.png" pageId="7" pageNumber="186">Fig. 4G</figureCitation>
). This could suggest that the specimens were able to extract the content of the diatom to digest it.
</paragraph>
</subSubSection>
</treatment>
</document>
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