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<document id="9DA8F1AD42E3ADA421C72EDA94EC106F" ID-CLB-Dataset="300095" ID-DOI="10.1007/s13127-023-00633-8" ID-GBIF-Dataset="fb16315d-7955-45b4-846b-63c23fe5b879" ID-ISSN="1618-1077" ID-Zenodo-Dep="12765007" IM.bibliography_approvedBy="julia" IM.illustrations_approvedBy="julia" IM.taxonomicNames_approvedBy="julia" IM.treatments_approvedBy="julia" checkinTime="1721248065677" checkinUser="felipe" docAuthor="Tilic, Ekin &amp; Rouse, Greg W." docDate="2024" docId="03CE1244FFE1FFF56F8EC9A05CAFFD2E" docLanguage="en" docName="OrgDivEvol.24.67-83.pdf" docOrigin="Organisms Diversity &amp; Evolution 24 (1)" docSource="http://dx.doi.org/10.1007/s13127-023-00633-8" docStyle="DocumentStyle:92260E183F2ECC5FAA0643175D3F67A1.1:OrgDivEvol.2010-.journal_article.open" docStyleId="92260E183F2ECC5FAA0643175D3F67A1" docStyleName="OrgDivEvol.2010-.journal_article.open" docStyleVersion="1" docTitle="Veneriserva Rossi 1984" docType="treatment" docVersion="2" lastPageNumber="74" masterDocId="FFF76A3CFFE5FFFD6F06CC2A5E41FFE9" masterDocTitle="Hardly Venuss servant-morphological adaptations of Veneriserva to an endoparasitic lifestyle and its phylogenetic position within Dorvilleidae (Annelida)" masterLastPageNumber="83" masterPageNumber="67" pageNumber="71" updateTime="1721754846505" updateUser="ExternalLinkService" zenodo-license-document="CC-BY-4.0">
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<mods:title id="9EEE60AA346E896B51D66965065A034C">Hardly Venuss servant-morphological adaptations of Veneriserva to an endoparasitic lifestyle and its phylogenetic position within Dorvilleidae (Annelida)</mods:title>
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<subSubSection id="C37DF0D9FFE1FFF96F8EC9A05C2FFA4C" box="[136,622,1418,1445]" pageId="4" pageNumber="71" type="nomenclature">
<paragraph id="8BD8A352FFE1FFF96F8EC9A05C2FFA4C" blockId="4.[136,622,1418,1445]" box="[136,622,1418,1445]" pageId="4" pageNumber="71">
<heading id="D090143EFFE1FFF96F8EC9A05C2FFA4C" box="[136,622,1418,1445]" fontSize="10" level="2" pageId="4" pageNumber="71" reason="9">
<emphasis id="B9137F40FFE1FFF96F8EC9A05C2FFA4C" bold="true" box="[136,622,1418,1445]" pageId="4" pageNumber="71">
Position of
<taxonomicName id="4C67D8D1FFE1FFF96E16C9A05FDDFA4C" ID-CoL="84VMM" authority="Rossi, 1984 " authorityName="Rossi" authorityYear="1984" box="[272,412,1418,1445]" class="Polychaeta" family="Dorvilleidae" genus="Veneriserva" kingdom="Animalia" order="Eunicida" pageId="4" pageNumber="71" phylum="Annelida" rank="genus">
<emphasis id="B9137F40FFE1FFF96E16C9A05FDDFA4C" bold="true" box="[272,412,1418,1445]" italics="true" pageId="4" pageNumber="71">Veneriserva</emphasis>
</taxonomicName>
within
<emphasis id="B9137F40FFE1FFF96EF0C9A05C2FFA4C" bold="true" box="[502,622,1418,1445]" italics="true" pageId="4" pageNumber="71">Aphrodita</emphasis>
</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="C37DF0D9FFE1FFFB6F8EC9E55BEAF939" lastPageId="6" lastPageNumber="73" pageId="4" pageNumber="71" type="biology_ecology">
<paragraph id="8BD8A352FFE1FFF96F8EC9E55A3AFD4C" blockId="4.[136,773,1487,1911]" lastBlockId="4.[816,1452,153,944]" pageId="4" pageNumber="71">
The µCT scan of a parasitized
<taxonomicName id="4C67D8D1FFE1FFF96ED0C9E55C2AFA0E" authorityName="Essenberg" authorityYear="1917" box="[470,619,1487,1511]" class="Polychaeta" family="Aphroditidae" genus="Aphrodita" kingdom="Animalia" order="Phyllodocida" pageId="4" pageNumber="71" phylum="Annelida" rank="species" species="longipalpa">
<emphasis id="B9137F40FFE1FFF96ED0C9E55C2AFA0E" box="[470,619,1487,1511]" italics="true" pageId="4" pageNumber="71">A. longipalpa</emphasis>
</taxonomicName>
allowed visualizing in 3D the position of
<taxonomicName id="4C67D8D1FFE1FFF96ECFC9DB5C0DF9E1" box="[457,588,1521,1544]" class="Polychaeta" family="Dorvilleidae" genus="Veneriserva" kingdom="Animalia" order="Eunicida" pageId="4" pageNumber="71" phylum="Annelida" rank="genus">
<emphasis id="B9137F40FFE1FFF96ECFC9DB5C0DF9E1" box="[457,588,1521,1544]" italics="true" pageId="4" pageNumber="71">Veneriserva</emphasis>
</taxonomicName>
inside their host without dissection and in their natural condition (
<figureCitation id="135CBFD7FFE1FFF96DBDCA3B5CB2F9C0" box="[699,755,1553,1577]" captionStart="Fig" captionStartId="6.[137,166,1429,1450]" captionTargetBox="[165,1423,154,1382]" captionTargetId="figure-145@6.[165,1423,154,1382]" captionTargetPageId="6" captionText="Fig.3 µCT visualization of parasites within Aphrodita longipalpa. A 3D rendering of parasites shown within the projection of the host body. BD Virtual dissections of surface renderings, showing crosssections of the host across three consecutive body regions, from anterior to posterior. Raw image data from the micro-CT stack, illustrating a horizontal section through the host (E) and a sagittal section (F). Head of the juvenile parasite is magnified to display the prominent jaws in white. Abbreviations—ja jaws, ne nephridia, pha pharynx. Female Veneriserva pygoclava is shown in yellow or with yellow arrowheads and the juvenile V. pygoclava in blue or with blue arrowheads" figureDoi="http://doi.org/10.5281/zenodo.12765013" httpUri="https://zenodo.org/record/12765013/files/figure.png" pageId="4" pageNumber="71">Fig. 3</figureCitation>
). In the
<taxonomicName id="4C67D8D1FFE1FFF96FD4CA195F01F9A2" authorityName="Linnaeus" authorityYear="1758" box="[210,320,1587,1611]" class="Polychaeta" family="Aphroditidae" genus="Aphrodita" kingdom="Animalia" order="Phyllodocida" pageId="4" pageNumber="71" phylum="Annelida" rank="genus">
<emphasis id="B9137F40FFE1FFF96FD4CA195F01F9A2" box="[210,320,1587,1611]" italics="true" pageId="4" pageNumber="71">Aphrodita</emphasis>
</taxonomicName>
specimen scanned, a single juvenile and a single female
<taxonomicName id="4C67D8D1FFE1FFF96E30CA7F5F80F985" box="[310,449,1620,1644]" class="Polychaeta" family="Dorvilleidae" genus="Veneriserva" kingdom="Animalia" order="Eunicida" pageId="4" pageNumber="71" phylum="Annelida" rank="species" species="pygoclava">
<emphasis id="B9137F40FFE1FFF96E30CA7F5F80F985" box="[310,449,1620,1644]" italics="true" pageId="4" pageNumber="71">V. pygoclava</emphasis>
</taxonomicName>
were present. The juvenile is likely to be a male, as
<specimenCount id="9D6168DBFFE1FFF96E8BCA5C5C52F964" box="[397,531,1653,1677]" count="2" pageId="4" pageNumber="71" type="female">two females</specimenCount>
were never observed together in a single host.
<taxonomicName id="4C67D8D1FFE1FFF96EAACABD5CEDF946" box="[428,684,1687,1711]" class="Polychaeta" family="Dorvilleidae" genus="Veneriserva" kingdom="Animalia" order="Eunicida" pageId="4" pageNumber="71" phylum="Annelida" rank="species" species="pygoclava">
<emphasis id="B9137F40FFE1FFF96EAACABD5CEDF946" box="[428,684,1687,1711]" italics="true" pageId="4" pageNumber="71">Veneriserva pygoclava</emphasis>
</taxonomicName>
resided within the coelomic cavity of the host, largely occupying the lateral and ventral coelomic spaces (
<figureCitation id="135CBFD7FFE1FFF96D41CAF35C3DF918" box="[583,636,1753,1777]" captionStart="Fig" captionStartId="6.[137,166,1429,1450]" captionTargetBox="[165,1423,154,1382]" captionTargetId="figure-145@6.[165,1423,154,1382]" captionTargetPageId="6" captionText="Fig.3 µCT visualization of parasites within Aphrodita longipalpa. A 3D rendering of parasites shown within the projection of the host body. BD Virtual dissections of surface renderings, showing crosssections of the host across three consecutive body regions, from anterior to posterior. Raw image data from the micro-CT stack, illustrating a horizontal section through the host (E) and a sagittal section (F). Head of the juvenile parasite is magnified to display the prominent jaws in white. Abbreviations—ja jaws, ne nephridia, pha pharynx. Female Veneriserva pygoclava is shown in yellow or with yellow arrowheads and the juvenile V. pygoclava in blue or with blue arrowheads" figureDoi="http://doi.org/10.5281/zenodo.12765013" httpUri="https://zenodo.org/record/12765013/files/figure.png" pageId="4" pageNumber="71">Fig. 3</figureCitation>
). The larger female was approximately
<quantity id="4C9F0EB7FFE1FFF96EACCAD15C4BF8FB" box="[426,522,1787,1811]" metricMagnitude="-2" metricUnit="m" metricValue="7.709999999999999" pageId="4" pageNumber="71" unit="mm" value="77.1">77.1 mm</quantity>
long, 10 times the size of the juvenile (±
<quantity id="4C9F0EB7FFE1FFF96E54CB365FEBF8DD" box="[338,426,1820,1844]" metricMagnitude="-3" metricUnit="m" metricValue="7.2" pageId="4" pageNumber="71" unit="mm" value="7.2">7.2 mm</quantity>
), and about twice the length of the host (±
<quantity id="4C9F0EB7FFE1FFF96E04CB175F22F8BC" box="[258,355,1853,1877]" metricMagnitude="-2" metricUnit="m" metricValue="4.27" pageId="4" pageNumber="71" unit="mm" value="42.7">42.7 mm</quantity>
) (
<figureCitation id="135CBFD7FFE1FFF96E85CB175F91F8BC" box="[387,464,1853,1877]" captionStart="Fig" captionStartId="6.[137,166,1429,1450]" captionTargetBox="[165,1423,154,1382]" captionTargetId="figure-145@6.[165,1423,154,1382]" captionTargetPageId="6" captionText="Fig.3 µCT visualization of parasites within Aphrodita longipalpa. A 3D rendering of parasites shown within the projection of the host body. BD Virtual dissections of surface renderings, showing crosssections of the host across three consecutive body regions, from anterior to posterior. Raw image data from the micro-CT stack, illustrating a horizontal section through the host (E) and a sagittal section (F). Head of the juvenile parasite is magnified to display the prominent jaws in white. Abbreviations—ja jaws, ne nephridia, pha pharynx. Female Veneriserva pygoclava is shown in yellow or with yellow arrowheads and the juvenile V. pygoclava in blue or with blue arrowheads" figureDoi="http://doi.org/10.5281/zenodo.12765013" httpUri="https://zenodo.org/record/12765013/files/figure.png" pageId="4" pageNumber="71">Fig. 3A</figureCitation>
). The female was positioned in a U-shaped coil with the posterior and anterior ends both located near the anterior of the host. The juvenile was located near the anterior end of the female (
<figureCitation id="135CBFD7FFE1FFF96A1CCC915D02FF1D" captionStart="Fig" captionStartId="6.[137,166,1429,1450]" captionTargetBox="[165,1423,154,1382]" captionTargetId="figure-145@6.[165,1423,154,1382]" captionTargetPageId="6" captionText="Fig.3 µCT visualization of parasites within Aphrodita longipalpa. A 3D rendering of parasites shown within the projection of the host body. BD Virtual dissections of surface renderings, showing crosssections of the host across three consecutive body regions, from anterior to posterior. Raw image data from the micro-CT stack, illustrating a horizontal section through the host (E) and a sagittal section (F). Head of the juvenile parasite is magnified to display the prominent jaws in white. Abbreviations—ja jaws, ne nephridia, pha pharynx. Female Veneriserva pygoclava is shown in yellow or with yellow arrowheads and the juvenile V. pygoclava in blue or with blue arrowheads" figureDoi="http://doi.org/10.5281/zenodo.12765013" httpUri="https://zenodo.org/record/12765013/files/figure.png" pageId="4" pageNumber="71">Fig. 3A, B, E, F</figureCitation>
). Other female
<taxonomicName id="4C67D8D1FFE1FFF96CF6CCF75A32FF1D" box="[1008,1139,221,244]" class="Polychaeta" family="Dorvilleidae" genus="Veneriserva" kingdom="Animalia" order="Eunicida" pageId="4" pageNumber="71" phylum="Annelida" rank="genus">
<emphasis id="B9137F40FFE1FFF96CF6CCF75A32FF1D" box="[1008,1139,221,244]" italics="true" pageId="4" pageNumber="71">Veneriserva</emphasis>
</taxonomicName>
with an even larger parasite to host body length ratio were observed in some of the dissected specimens (
<figureCitation id="135CBFD7FFE1FFF96B04CD355A2EFEDE" box="[1026,1135,287,311]" captionStart="◂" captionStartId="4.[122,136,145,168]" captionTargetId="figure-13@3.[147,1440,154,1854]" captionTargetPageId="3" captionText="◂Fig. 1 Live photos and dissection of parasitized Aphrodita longipalpa and Veneriserva pygoclava. A Ventral view of A. longipalpa. B Dorsal view of A. longipalpa with removed feltage chaetae, revealing the parasite visible through the body wall. C Ventrally dissected A. longipalpa, exposing the sizable female parasite. Veneriserva pygoclava individuals within the host are indicated by arrowheads. D Juvenile female V. pygoclava, with developing oocytes visible through the body wall along the mid-dorsal orange line. E Female V. pygoclava showing the mid-dorsal orange pigmentation and the white mark at the base of the prostomium. F Male V. pygoclava. G A large female and smaller male V. pygoclava, extracted from the same host. The pygidium is club-shaped in both males and females and juveniles. H Juvenile V. pygoclava shown from multiple angles, characterized by a complete white coloration; black jaws are magnified in panel" figureDoi="http://doi.org/10.5281/zenodo.12802034" httpUri="https://zenodo.org/record/12802034/files/figure.png" pageId="4" pageNumber="71">Fig. 1C, G</figureCitation>
). In these, the parasites could be observed making multiple coils within the ventral coelomic space, and also extending dorsally (
<figureCitation id="135CBFD7FFE1FFF96BE8CD4B5B19FE93" box="[1262,1368,353,378]" captionStart="◂" captionStartId="4.[122,136,145,168]" captionTargetId="figure-13@3.[147,1440,154,1854]" captionTargetPageId="3" captionText="◂Fig. 1 Live photos and dissection of parasitized Aphrodita longipalpa and Veneriserva pygoclava. A Ventral view of A. longipalpa. B Dorsal view of A. longipalpa with removed feltage chaetae, revealing the parasite visible through the body wall. C Ventrally dissected A. longipalpa, exposing the sizable female parasite. Veneriserva pygoclava individuals within the host are indicated by arrowheads. D Juvenile female V. pygoclava, with developing oocytes visible through the body wall along the mid-dorsal orange line. E Female V. pygoclava showing the mid-dorsal orange pigmentation and the white mark at the base of the prostomium. F Male V. pygoclava. G A large female and smaller male V. pygoclava, extracted from the same host. The pygidium is club-shaped in both males and females and juveniles. H Juvenile V. pygoclava shown from multiple angles, characterized by a complete white coloration; black jaws are magnified in panel" figureDoi="http://doi.org/10.5281/zenodo.12802034" httpUri="https://zenodo.org/record/12802034/files/figure.png" pageId="4" pageNumber="71">Fig. 1AC</figureCitation>
), taking up a very large area within the hosts body. Neither the musculature nor the gut of the host appeared to be substantially damaged by the parasite (
<figureCitation id="135CBFD7FFE1FFF96B4FCDEF5AF7FE37" box="[1097,1206,453,478]" captionStart="Fig" captionStartId="6.[137,166,1429,1450]" captionTargetBox="[165,1423,154,1382]" captionTargetId="figure-145@6.[165,1423,154,1382]" captionTargetPageId="6" captionText="Fig.3 µCT visualization of parasites within Aphrodita longipalpa. A 3D rendering of parasites shown within the projection of the host body. BD Virtual dissections of surface renderings, showing crosssections of the host across three consecutive body regions, from anterior to posterior. Raw image data from the micro-CT stack, illustrating a horizontal section through the host (E) and a sagittal section (F). Head of the juvenile parasite is magnified to display the prominent jaws in white. Abbreviations—ja jaws, ne nephridia, pha pharynx. Female Veneriserva pygoclava is shown in yellow or with yellow arrowheads and the juvenile V. pygoclava in blue or with blue arrowheads" figureDoi="http://doi.org/10.5281/zenodo.12765013" httpUri="https://zenodo.org/record/12765013/files/figure.png" pageId="4" pageNumber="71">Fig. 3BD</figureCitation>
). Despite the relatively large size of many of the
<taxonomicName id="4C67D8D1FFE1FFF96B3CCDCD5A8AFE16" authorityName="Essenberg" authorityYear="1917" box="[1082,1227,487,511]" class="Polychaeta" family="Aphroditidae" genus="Aphrodita" kingdom="Animalia" order="Phyllodocida" pageId="4" pageNumber="71" phylum="Annelida" rank="species" species="longipalpa">
<emphasis id="B9137F40FFE1FFF96B3CCDCD5A8AFE16" box="[1082,1227,487,511]" italics="true" pageId="4" pageNumber="71">A. longipalpa</emphasis>
</taxonomicName>
collected, no gonads or gametes of any stage were observed in any sampled hosts, infected or not. Specimens were observed from the months December, March, May, July, September, and October over the period of 2017 to 2023; so, the breeding season may be over the winter/spring months.
</paragraph>
<paragraph id="8BD8A352FFE1FFF96C49CE855AF1FC59" blockId="4.[816,1452,153,944]" pageId="4" pageNumber="71">
Interestingly, the
<specimenCount id="9D6168DBFFE1FFF96B08CE855AD4FD2E" box="[1038,1173,687,711]" count="2" pageId="4" pageNumber="71" type="juvenile">two juvenile</specimenCount>
<taxonomicName id="4C67D8D1FFE1FFF96B98CE855B72FD2E" authorityName="Essenberg" authorityYear="1917" box="[1182,1331,687,711]" class="Polychaeta" family="Aphroditidae" genus="Aphrodita" kingdom="Animalia" order="Phyllodocida" pageId="4" pageNumber="71" phylum="Annelida" rank="species" species="longipalpa">
<emphasis id="B9137F40FFE1FFF96B98CE855B72FD2E" box="[1182,1331,687,711]" italics="true" pageId="4" pageNumber="71">A. longipalpa</emphasis>
</taxonomicName>
specimens examined (both less than
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in length) did not show any signs of infection upon external inspection of the ventral side. However, histological sectioning of these specimens revealed a single juvenile
<taxonomicName id="4C67D8D1FFE1FFF96B45CF1F5A8DFCA5" box="[1091,1228,820,844]" class="Polychaeta" family="Dorvilleidae" genus="Veneriserva" kingdom="Animalia" order="Eunicida" pageId="4" pageNumber="71" phylum="Annelida" rank="species" species="pygoclava">
<emphasis id="B9137F40FFE1FFF96B45CF1F5A8DFCA5" box="[1091,1228,820,844]" italics="true" pageId="4" pageNumber="71">V. pygoclava</emphasis>
</taxonomicName>
in both (
<figureCitation id="135CBFD7FFE1FFF96A32CF1E5BDFFCA5" box="[1332,1438,820,844]" captionStart="Fig" captionStartId="7.[136,165,1303,1324]" captionTargetBox="[144,1444,154,1255]" captionTargetId="figure-239@7.[144,1444,154,1255]" captionTargetPageId="7" captionText="Fig. 4 AZAN-stained paraffin histology. A Histological cross-section of a juvenile Aphrodita longipalpa featuring an endoparasitic immature Veneriserva pygoclava (denoted by a star). B Longitudinal section of V. pygoclava, highlighting the absence of a through gut, and continuous uninterrupted mesenteries. CH Cross-sections through the anterior region of V. pygoclava, showing the muscularized pharynx with jaws culminating in blind termination at section G. Abbreviations—ac acicula, br brain, df dorsal felt, el elytra, ja jaws, mo mouth, ne nephridium, pha pharynx, vnc ventral nerve cord" figureDoi="http://doi.org/10.5281/zenodo.12765015" httpUri="https://zenodo.org/record/12765015/files/figure.png" pageId="4" pageNumber="71">Fig. 4A, B</figureCitation>
). These
<taxonomicName id="4C67D8D1FFE1FFF96C7CCF7C5A49FC84" box="[890,1032,853,877]" class="Polychaeta" family="Dorvilleidae" genus="Veneriserva" kingdom="Animalia" order="Eunicida" pageId="4" pageNumber="71" phylum="Annelida" rank="species" species="pygoclava">
<emphasis id="B9137F40FFE1FFF96C7CCF7C5A49FC84" box="[890,1032,853,877]" italics="true" pageId="4" pageNumber="71">V. pygoclava</emphasis>
</taxonomicName>
specimens were juveniles themselves and did not show any signs of gametogenesis (
<figureCitation id="135CBFD7FFE1FFF96A33CF5D5B3FFC66" box="[1333,1406,887,911]" captionStart="Fig" captionStartId="7.[136,165,1303,1324]" captionTargetBox="[144,1444,154,1255]" captionTargetId="figure-239@7.[144,1444,154,1255]" captionTargetPageId="7" captionText="Fig. 4 AZAN-stained paraffin histology. A Histological cross-section of a juvenile Aphrodita longipalpa featuring an endoparasitic immature Veneriserva pygoclava (denoted by a star). B Longitudinal section of V. pygoclava, highlighting the absence of a through gut, and continuous uninterrupted mesenteries. CH Cross-sections through the anterior region of V. pygoclava, showing the muscularized pharynx with jaws culminating in blind termination at section G. Abbreviations—ac acicula, br brain, df dorsal felt, el elytra, ja jaws, mo mouth, ne nephridium, pha pharynx, vnc ventral nerve cord" figureDoi="http://doi.org/10.5281/zenodo.12765015" httpUri="https://zenodo.org/record/12765015/files/figure.png" pageId="4" pageNumber="71">Fig. 4B</figureCitation>
), so sex determination was not possible.
</paragraph>
<paragraph id="8BD8A352FFE1FFF96C36CFF35DEEFC1D" blockId="4.[816,943,985,1012]" box="[816,943,985,1012]" pageId="4" pageNumber="71">
<heading id="D090143EFFE1FFF96C36CFF35DEEFC1D" bold="true" box="[816,943,985,1012]" fontSize="11" level="1" pageId="4" pageNumber="71" reason="1">
<emphasis id="B9137F40FFE1FFF96C36CFF35DEEFC1D" bold="true" box="[816,943,985,1012]" pageId="4" pageNumber="71">Oogenesis</emphasis>
</heading>
</paragraph>
<paragraph id="8BD8A352FFE1FFF96C36C8375AE3F89E" blockId="4.[816,1453,1053,1911]" pageId="4" pageNumber="71">
Oogenesis in
<taxonomicName id="4C67D8D1FFE1FFF96CC5C8345AFBFBDC" box="[963,1210,1053,1077]" class="Polychaeta" family="Dorvilleidae" genus="Veneriserva" kingdom="Animalia" order="Eunicida" pageId="4" pageNumber="71" phylum="Annelida" rank="species" species="pygoclava">
<emphasis id="B9137F40FFE1FFF96CC5C8345AFBFBDC" box="[963,1210,1053,1077]" italics="true" pageId="4" pageNumber="71">Veneriserva pygoclava</emphasis>
</taxonomicName>
occurs in segmentally repeated ovaries. The oogonia of
<taxonomicName id="4C67D8D1FFE1FFF96B9BC86A5B69FBBE" box="[1181,1320,1087,1111]" class="Polychaeta" family="Dorvilleidae" genus="Veneriserva" kingdom="Animalia" order="Eunicida" pageId="4" pageNumber="71" phylum="Annelida" rank="species" species="pygoclava">
<emphasis id="B9137F40FFE1FFF96B9BC86A5B69FBBE" box="[1181,1320,1087,1111]" italics="true" pageId="4" pageNumber="71">V. pygoclava</emphasis>
</taxonomicName>
proliferated from the ventral surface of the dorsal blood vessel (
<figureCitation id="135CBFD7FFE1FFF96A5BC84A5D02FB73" captionStart="◂" captionStartId="10.[122,136,145,168]" captionTargetId="figure-13@9.[147,1440,154,1854]" captionTargetPageId="9" captionText="◂Fig.5 Gametogenesis in male and female Veneriserva pygoclava. AD Semi-thin histological sections of female Veneriserva pygoclava, stained with toluidine blue. A Cross-section of a female Veneriserva. B Close-up of large mature oocytes without discernible nurse cells. C Developing oocytes attached to mesenteries (mes), and oogonia proliferating from the ventral side of the dorsal blood vessel (bv). D Details of vitellogenic oocytes and nurse cells. Arrowheads indicate brownstained yolk platelets and yolk bodies. E Live sperm cells captured in a light micrograph. FG Cross-sections of male Veneriserva. Note the absence of a gut in the cross-sections. Abbreviations—ac acicula, acr acrosome, bv blood vessel, coe coelomic cavity, mes mesentery, nc nurse cell, nn nurse cell nucleus, nu sperm cell nucleus, Oo oocyte, on oocyte nucleus, sp spermatogonia, vnc ventral nerve cord" figureDoi="http://doi.org/10.5281/zenodo.12802038" httpUri="https://zenodo.org/record/12802038/files/figure.png" pageId="4" pageNumber="71">Fig. 5A, C</figureCitation>
) and the gonads were attached to the intersegmental septa (mesenteries) (
<figureCitation id="135CBFD7FFE1FFF96B1CC8895A24FB52" box="[1050,1125,1187,1211]" captionStart="◂" captionStartId="10.[122,136,145,168]" captionTargetId="figure-13@9.[147,1440,154,1854]" captionTargetPageId="9" captionText="◂Fig.5 Gametogenesis in male and female Veneriserva pygoclava. AD Semi-thin histological sections of female Veneriserva pygoclava, stained with toluidine blue. A Cross-section of a female Veneriserva. B Close-up of large mature oocytes without discernible nurse cells. C Developing oocytes attached to mesenteries (mes), and oogonia proliferating from the ventral side of the dorsal blood vessel (bv). D Details of vitellogenic oocytes and nurse cells. Arrowheads indicate brownstained yolk platelets and yolk bodies. E Live sperm cells captured in a light micrograph. FG Cross-sections of male Veneriserva. Note the absence of a gut in the cross-sections. Abbreviations—ac acicula, acr acrosome, bv blood vessel, coe coelomic cavity, mes mesentery, nc nurse cell, nn nurse cell nucleus, nu sperm cell nucleus, Oo oocyte, on oocyte nucleus, sp spermatogonia, vnc ventral nerve cord" figureDoi="http://doi.org/10.5281/zenodo.12802038" httpUri="https://zenodo.org/record/12802038/files/figure.png" pageId="4" pageNumber="71">Fig. 5C</figureCitation>
). Each developing oocyte was directly connected to a nurse cell by intercellular cytoplasmic bridges (
<figureCitation id="135CBFD7FFE1FFF96CC4C8CF5A4CFB17" box="[962,1037,1253,1278]" captionStart="◂" captionStartId="10.[122,136,145,168]" captionTargetId="figure-13@9.[147,1440,154,1854]" captionTargetPageId="9" captionText="◂Fig.5 Gametogenesis in male and female Veneriserva pygoclava. AD Semi-thin histological sections of female Veneriserva pygoclava, stained with toluidine blue. A Cross-section of a female Veneriserva. B Close-up of large mature oocytes without discernible nurse cells. C Developing oocytes attached to mesenteries (mes), and oogonia proliferating from the ventral side of the dorsal blood vessel (bv). D Details of vitellogenic oocytes and nurse cells. Arrowheads indicate brownstained yolk platelets and yolk bodies. E Live sperm cells captured in a light micrograph. FG Cross-sections of male Veneriserva. Note the absence of a gut in the cross-sections. Abbreviations—ac acicula, acr acrosome, bv blood vessel, coe coelomic cavity, mes mesentery, nc nurse cell, nn nurse cell nucleus, nu sperm cell nucleus, Oo oocyte, on oocyte nucleus, sp spermatogonia, vnc ventral nerve cord" figureDoi="http://doi.org/10.5281/zenodo.12802038" httpUri="https://zenodo.org/record/12802038/files/figure.png" pageId="4" pageNumber="71">Fig. 5D</figureCitation>
). The nucleus of the oocyte appeared to undergo numerous morphological changes during oogenesis, as it appeared heterochromatic in early stages and enlarged and became euchromatic with a single prominent nucleolus during the vitellogenic phase (
<figureCitation id="135CBFD7FFE1FFF96B83C9415AB3FA6A" box="[1157,1266,1387,1411]" captionStart="◂" captionStartId="10.[122,136,145,168]" captionTargetId="figure-13@9.[147,1440,154,1854]" captionTargetPageId="9" captionText="◂Fig.5 Gametogenesis in male and female Veneriserva pygoclava. AD Semi-thin histological sections of female Veneriserva pygoclava, stained with toluidine blue. A Cross-section of a female Veneriserva. B Close-up of large mature oocytes without discernible nurse cells. C Developing oocytes attached to mesenteries (mes), and oogonia proliferating from the ventral side of the dorsal blood vessel (bv). D Details of vitellogenic oocytes and nurse cells. Arrowheads indicate brownstained yolk platelets and yolk bodies. E Live sperm cells captured in a light micrograph. FG Cross-sections of male Veneriserva. Note the absence of a gut in the cross-sections. Abbreviations—ac acicula, acr acrosome, bv blood vessel, coe coelomic cavity, mes mesentery, nc nurse cell, nn nurse cell nucleus, nu sperm cell nucleus, Oo oocyte, on oocyte nucleus, sp spermatogonia, vnc ventral nerve cord" figureDoi="http://doi.org/10.5281/zenodo.12802038" httpUri="https://zenodo.org/record/12802038/files/figure.png" pageId="4" pageNumber="71">Fig. 5B, C</figureCitation>
). The nurse cells appeared to undergo a striking morphological change after the onset of vitellogenesis, making them easy to distinguish from their neighboring oocytes. During vitellogenesis, the volume of the nurse cell nucleus rapidly increased and was always dense and heterochromatic, whereas the nuclei of the oocytes were all euchromatic at this stage (
<figureCitation id="135CBFD7FFE1FFF96A57CA195BDFF9A2" box="[1361,1438,1587,1611]" captionStart="◂" captionStartId="10.[122,136,145,168]" captionTargetId="figure-13@9.[147,1440,154,1854]" captionTargetPageId="9" captionText="◂Fig.5 Gametogenesis in male and female Veneriserva pygoclava. AD Semi-thin histological sections of female Veneriserva pygoclava, stained with toluidine blue. A Cross-section of a female Veneriserva. B Close-up of large mature oocytes without discernible nurse cells. C Developing oocytes attached to mesenteries (mes), and oogonia proliferating from the ventral side of the dorsal blood vessel (bv). D Details of vitellogenic oocytes and nurse cells. Arrowheads indicate brownstained yolk platelets and yolk bodies. E Live sperm cells captured in a light micrograph. FG Cross-sections of male Veneriserva. Note the absence of a gut in the cross-sections. Abbreviations—ac acicula, acr acrosome, bv blood vessel, coe coelomic cavity, mes mesentery, nc nurse cell, nn nurse cell nucleus, nu sperm cell nucleus, Oo oocyte, on oocyte nucleus, sp spermatogonia, vnc ventral nerve cord" figureDoi="http://doi.org/10.5281/zenodo.12802038" httpUri="https://zenodo.org/record/12802038/files/figure.png" pageId="4" pageNumber="71">Fig. 5C</figureCitation>
). Another difference between vitellogenic oocytes and nurse cells was the size and distribution of yolk platelets. Nurse cells never contained ripe yolk bodies, but only small-sized yolk platelets arranged in clusters (
<figureCitation id="135CBFD7FFE1FFF96BB7CA925ABFF939" box="[1201,1278,1720,1744]" captionStart="◂" captionStartId="10.[122,136,145,168]" captionTargetId="figure-13@9.[147,1440,154,1854]" captionTargetPageId="9" captionText="◂Fig.5 Gametogenesis in male and female Veneriserva pygoclava. AD Semi-thin histological sections of female Veneriserva pygoclava, stained with toluidine blue. A Cross-section of a female Veneriserva. B Close-up of large mature oocytes without discernible nurse cells. C Developing oocytes attached to mesenteries (mes), and oogonia proliferating from the ventral side of the dorsal blood vessel (bv). D Details of vitellogenic oocytes and nurse cells. Arrowheads indicate brownstained yolk platelets and yolk bodies. E Live sperm cells captured in a light micrograph. FG Cross-sections of male Veneriserva. Note the absence of a gut in the cross-sections. Abbreviations—ac acicula, acr acrosome, bv blood vessel, coe coelomic cavity, mes mesentery, nc nurse cell, nn nurse cell nucleus, nu sperm cell nucleus, Oo oocyte, on oocyte nucleus, sp spermatogonia, vnc ventral nerve cord" figureDoi="http://doi.org/10.5281/zenodo.12802038" httpUri="https://zenodo.org/record/12802038/files/figure.png" pageId="4" pageNumber="71">Fig. 5D</figureCitation>
). While oocytes kept growing reaching a final diameter of about 100 µm, nurse cells had already reached their final diameter of about 30 µm during vitellogenesis. The nurse cells then appear to undergo a decrease in diameter until they are finally incorporated into the oocytes (
<figureCitation id="135CBFD7FFE1FFF96B4CCB755AD4F89E" box="[1098,1173,1887,1911]" captionStart="◂" captionStartId="10.[122,136,145,168]" captionTargetId="figure-13@9.[147,1440,154,1854]" captionTargetPageId="9" captionText="◂Fig.5 Gametogenesis in male and female Veneriserva pygoclava. AD Semi-thin histological sections of female Veneriserva pygoclava, stained with toluidine blue. A Cross-section of a female Veneriserva. B Close-up of large mature oocytes without discernible nurse cells. C Developing oocytes attached to mesenteries (mes), and oogonia proliferating from the ventral side of the dorsal blood vessel (bv). D Details of vitellogenic oocytes and nurse cells. Arrowheads indicate brownstained yolk platelets and yolk bodies. E Live sperm cells captured in a light micrograph. FG Cross-sections of male Veneriserva. Note the absence of a gut in the cross-sections. Abbreviations—ac acicula, acr acrosome, bv blood vessel, coe coelomic cavity, mes mesentery, nc nurse cell, nn nurse cell nucleus, nu sperm cell nucleus, Oo oocyte, on oocyte nucleus, sp spermatogonia, vnc ventral nerve cord" figureDoi="http://doi.org/10.5281/zenodo.12802038" httpUri="https://zenodo.org/record/12802038/files/figure.png" pageId="4" pageNumber="71">Fig. 5B</figureCitation>
).
</paragraph>
<caption id="DF18F3DAFFE0FFF86F8FC9A55AFBFA1D" ID-DOI="http://doi.org/10.5281/zenodo.12765011" ID-Zenodo-Dep="12765011" httpUri="https://zenodo.org/record/12765011/files/figure.png" pageId="5" pageNumber="72" startId="5.[137,166,1423,1444]" subCaptionStartIDs="5.[816,916,1449,1471]" subCaptionStarts="figu" targetBox="[157,1430,155,1375]" targetPageId="5" targetType="figure">
<paragraph id="8BD8A352FFE0FFF86F8FC9A55AFBFA1D" blockId="5.[136,772,1422,1524]" lastBlockId="5.[816,1452,1422,1524]" pageId="5" pageNumber="72">
<emphasis id="B9137F40FFE0FFF86F8FC9A55EF9FA4D" bold="true" box="[137,184,1423,1444]" pageId="5" pageNumber="72">Fig. 2</emphasis>
Parasite abundance and distribution statistics. A total of 58
<taxonomicName id="4C67D8D1FFE0FFF86F8EC9835F0CFA57" authorityName="Essenberg" authorityYear="1917" box="[136,333,1449,1470]" class="Polychaeta" family="Aphroditidae" genus="Aphrodita" kingdom="Animalia" order="Phyllodocida" pageId="5" pageNumber="72" phylum="Annelida" rank="species" species="longipalpa">
<emphasis id="B9137F40FFE0FFF86F8EC9835F0CFA57" box="[136,333,1449,1470]" italics="true" pageId="5" pageNumber="72">Aphrodita longipalpa</emphasis>
</taxonomicName>
were dissected and examined for parasite presence. The upper horizontal bars graphically depict the proportional parasitism rates and the corresponding distribution among male, female, and juvenile parasites, along with various cohabitation configurations. The box plots show the relationship between host size and the occurrence of parasites, presented collectively and then individually for female, male, and juvenile parasites
</paragraph>
</caption>
<paragraph id="8BD8A352FFE0FFF86F8ECA195F0AF9A7" blockId="5.[136,331,1587,1614]" box="[136,331,1587,1614]" pageId="5" pageNumber="72">
<heading id="D090143EFFE0FFF86F8ECA195F0AF9A7" bold="true" box="[136,331,1587,1614]" fontSize="11" level="1" pageId="5" pageNumber="72" reason="1">
<emphasis id="B9137F40FFE0FFF86F8ECA195F0AF9A7" bold="true" box="[136,331,1587,1614]" pageId="5" pageNumber="72">Spermiogenesis</emphasis>
</heading>
</paragraph>
<paragraph id="8BD8A352FFE0FFF86F8ECA525B3EF985" blockId="5.[136,771,1655,1913]" lastBlockId="5.[816,1451,1587,1644]" pageId="5" pageNumber="72">
In the histologically sectioned male specimen, spermiogenesis occurred along the peritoneal lining (
<figureCitation id="135CBFD7FFE0FFF86DA0CAB35CB7F958" box="[678,758,1689,1713]" captionStart="◂" captionStartId="10.[122,136,145,168]" captionTargetId="figure-13@9.[147,1440,154,1854]" captionTargetPageId="9" captionText="◂Fig.5 Gametogenesis in male and female Veneriserva pygoclava. AD Semi-thin histological sections of female Veneriserva pygoclava, stained with toluidine blue. A Cross-section of a female Veneriserva. B Close-up of large mature oocytes without discernible nurse cells. C Developing oocytes attached to mesenteries (mes), and oogonia proliferating from the ventral side of the dorsal blood vessel (bv). D Details of vitellogenic oocytes and nurse cells. Arrowheads indicate brownstained yolk platelets and yolk bodies. E Live sperm cells captured in a light micrograph. FG Cross-sections of male Veneriserva. Note the absence of a gut in the cross-sections. Abbreviations—ac acicula, acr acrosome, bv blood vessel, coe coelomic cavity, mes mesentery, nc nurse cell, nn nurse cell nucleus, nu sperm cell nucleus, Oo oocyte, on oocyte nucleus, sp spermatogonia, vnc ventral nerve cord" figureDoi="http://doi.org/10.5281/zenodo.12802038" httpUri="https://zenodo.org/record/12802038/files/figure.png" pageId="5" pageNumber="72">Fig. 5F</figureCitation>
). Spermatogonia were found ventrally associated with the coelomic lining, near the dorsal blood vessel (
<figureCitation id="135CBFD7FFE0FFF86DAFCAF15CB4F91A" box="[681,757,1755,1779]" captionStart="◂" captionStartId="10.[122,136,145,168]" captionTargetId="figure-13@9.[147,1440,154,1854]" captionTargetPageId="9" captionText="◂Fig.5 Gametogenesis in male and female Veneriserva pygoclava. AD Semi-thin histological sections of female Veneriserva pygoclava, stained with toluidine blue. A Cross-section of a female Veneriserva. B Close-up of large mature oocytes without discernible nurse cells. C Developing oocytes attached to mesenteries (mes), and oogonia proliferating from the ventral side of the dorsal blood vessel (bv). D Details of vitellogenic oocytes and nurse cells. Arrowheads indicate brownstained yolk platelets and yolk bodies. E Live sperm cells captured in a light micrograph. FG Cross-sections of male Veneriserva. Note the absence of a gut in the cross-sections. Abbreviations—ac acicula, acr acrosome, bv blood vessel, coe coelomic cavity, mes mesentery, nc nurse cell, nn nurse cell nucleus, nu sperm cell nucleus, Oo oocyte, on oocyte nucleus, sp spermatogonia, vnc ventral nerve cord" figureDoi="http://doi.org/10.5281/zenodo.12802038" httpUri="https://zenodo.org/record/12802038/files/figure.png" pageId="5" pageNumber="72">Fig. 5F</figureCitation>
). Spermatocytes and mature sperm were found released into the coelomic cavity. Upon dissection, liberated sperm cells could be observed under the light microscope. Mature sperm of
<taxonomicName id="4C67D8D1FFE0FFF86FF8CB4B5FD7F890" box="[254,406,1889,1913]" class="Polychaeta" family="Dorvilleidae" genus="Veneriserva" kingdom="Animalia" order="Eunicida" pageId="5" pageNumber="72" phylum="Annelida" rank="species" species="pygoclava">
<emphasis id="B9137F40FFE0FFF86FF8CB4B5FD7F890" box="[254,406,1889,1913]" italics="true" pageId="5" pageNumber="72">V. pygoclava</emphasis>
</taxonomicName>
had a large conical acrosome (~ 12 µm long) and a spherical nucleus (6 µm in diameter) with no evidence of an emergent flagellum (
<figureCitation id="135CBFD7FFE0FFF86A23CA7E5B33F985" box="[1317,1394,1620,1644]" captionStart="◂" captionStartId="10.[122,136,145,168]" captionTargetId="figure-13@9.[147,1440,154,1854]" captionTargetPageId="9" captionText="◂Fig.5 Gametogenesis in male and female Veneriserva pygoclava. AD Semi-thin histological sections of female Veneriserva pygoclava, stained with toluidine blue. A Cross-section of a female Veneriserva. B Close-up of large mature oocytes without discernible nurse cells. C Developing oocytes attached to mesenteries (mes), and oogonia proliferating from the ventral side of the dorsal blood vessel (bv). D Details of vitellogenic oocytes and nurse cells. Arrowheads indicate brownstained yolk platelets and yolk bodies. E Live sperm cells captured in a light micrograph. FG Cross-sections of male Veneriserva. Note the absence of a gut in the cross-sections. Abbreviations—ac acicula, acr acrosome, bv blood vessel, coe coelomic cavity, mes mesentery, nc nurse cell, nn nurse cell nucleus, nu sperm cell nucleus, Oo oocyte, on oocyte nucleus, sp spermatogonia, vnc ventral nerve cord" figureDoi="http://doi.org/10.5281/zenodo.12802038" httpUri="https://zenodo.org/record/12802038/files/figure.png" pageId="5" pageNumber="72">Fig. 5E</figureCitation>
).
</paragraph>
<paragraph id="8BD8A352FFE0FFF86C36CABF5B1CF959" blockId="5.[816,1373,1685,1712]" box="[816,1373,1685,1712]" pageId="5" pageNumber="72">
<heading id="D090143EFFE0FFF86C36CABF5B1CF959" bold="true" box="[816,1373,1685,1712]" fontSize="11" level="1" pageId="5" pageNumber="72" reason="1">
<emphasis id="B9137F40FFE0FFF86C36CABF5B1CF959" bold="true" box="[816,1373,1685,1712]" pageId="5" pageNumber="72">Pharyngeal apparatus and lack of an intestine</emphasis>
</heading>
</paragraph>
<paragraph id="8BD8A352FFE0FFFB6C36CAF35BEAF939" blockId="5.[816,1451,1753,1911]" lastBlockId="6.[816,1451,1653,1744]" lastPageId="6" lastPageNumber="73" pageId="5" pageNumber="72">
The alimentary tract of
<taxonomicName id="4C67D8D1FFE0FFF86B48CAF05AA5F918" box="[1102,1252,1753,1777]" class="Polychaeta" family="Dorvilleidae" genus="Veneriserva" kingdom="Animalia" order="Eunicida" pageId="5" pageNumber="72" phylum="Annelida" rank="species" species="pygoclava">
<emphasis id="B9137F40FFE0FFF86B48CAF05AA5F918" box="[1102,1252,1753,1777]" italics="true" pageId="5" pageNumber="72">V. pygoclava</emphasis>
</taxonomicName>
was found to be unique and aberrant among Annelida, in that it consisted of a functional muscular axial proboscis with a jaw apparatus but with no through-gut (
<figureCitation id="135CBFD7FFE0FFF86B92CB175AAAF8BC" box="[1172,1259,1853,1877]" captionStart="Fig" captionStartId="6.[137,166,1429,1450]" captionTargetBox="[165,1423,154,1382]" captionTargetId="figure-145@6.[165,1423,154,1382]" captionTargetPageId="6" captionText="Fig.3 µCT visualization of parasites within Aphrodita longipalpa. A 3D rendering of parasites shown within the projection of the host body. BD Virtual dissections of surface renderings, showing crosssections of the host across three consecutive body regions, from anterior to posterior. Raw image data from the micro-CT stack, illustrating a horizontal section through the host (E) and a sagittal section (F). Head of the juvenile parasite is magnified to display the prominent jaws in white. Abbreviations—ja jaws, ne nephridia, pha pharynx. Female Veneriserva pygoclava is shown in yellow or with yellow arrowheads and the juvenile V. pygoclava in blue or with blue arrowheads" figureDoi="http://doi.org/10.5281/zenodo.12765013" httpUri="https://zenodo.org/record/12765013/files/figure.png" pageId="5" pageNumber="72">Figs. 3E</figureCitation>
, and
<figureCitation id="135CBFD7FFE0FFF86A2ECB175B2CF8BC" box="[1320,1389,1853,1877]" captionStart="Fig" captionStartId="7.[136,165,1303,1324]" captionTargetBox="[144,1444,154,1255]" captionTargetId="figure-239@7.[144,1444,154,1255]" captionTargetPageId="7" captionText="Fig. 4 AZAN-stained paraffin histology. A Histological cross-section of a juvenile Aphrodita longipalpa featuring an endoparasitic immature Veneriserva pygoclava (denoted by a star). B Longitudinal section of V. pygoclava, highlighting the absence of a through gut, and continuous uninterrupted mesenteries. CH Cross-sections through the anterior region of V. pygoclava, showing the muscularized pharynx with jaws culminating in blind termination at section G. Abbreviations—ac acicula, br brain, df dorsal felt, el elytra, ja jaws, mo mouth, ne nephridium, pha pharynx, vnc ventral nerve cord" figureDoi="http://doi.org/10.5281/zenodo.12765015" httpUri="https://zenodo.org/record/12765015/files/figure.png" pageId="5" pageNumber="72">4BH</figureCitation>
). The pharynx ended blindly (
<figureCitation id="135CBFD7FFE0FFF86B40CB755AF4F89E" box="[1094,1205,1887,1911]" captionStart="Fig" captionStartId="7.[136,165,1303,1324]" captionTargetBox="[144,1444,154,1255]" captionTargetId="figure-239@7.[144,1444,154,1255]" captionTargetPageId="7" captionText="Fig. 4 AZAN-stained paraffin histology. A Histological cross-section of a juvenile Aphrodita longipalpa featuring an endoparasitic immature Veneriserva pygoclava (denoted by a star). B Longitudinal section of V. pygoclava, highlighting the absence of a through gut, and continuous uninterrupted mesenteries. CH Cross-sections through the anterior region of V. pygoclava, showing the muscularized pharynx with jaws culminating in blind termination at section G. Abbreviations—ac acicula, br brain, df dorsal felt, el elytra, ja jaws, mo mouth, ne nephridium, pha pharynx, vnc ventral nerve cord" figureDoi="http://doi.org/10.5281/zenodo.12765015" httpUri="https://zenodo.org/record/12765015/files/figure.png" pageId="5" pageNumber="72">Fig. 4G, H</figureCitation>
) leaving a completely hollow body cavity only occupied by developing gametes in mature specimens (
<figureCitation id="135CBFD7FFE3FFFB6E89CABD5C40F946" box="[399,513,1686,1711]" captionStart="◂" captionStartId="10.[122,136,145,168]" captionTargetId="figure-13@9.[147,1440,154,1854]" captionTargetPageId="9" captionText="◂Fig.5 Gametogenesis in male and female Veneriserva pygoclava. AD Semi-thin histological sections of female Veneriserva pygoclava, stained with toluidine blue. A Cross-section of a female Veneriserva. B Close-up of large mature oocytes without discernible nurse cells. C Developing oocytes attached to mesenteries (mes), and oogonia proliferating from the ventral side of the dorsal blood vessel (bv). D Details of vitellogenic oocytes and nurse cells. Arrowheads indicate brownstained yolk platelets and yolk bodies. E Live sperm cells captured in a light micrograph. FG Cross-sections of male Veneriserva. Note the absence of a gut in the cross-sections. Abbreviations—ac acicula, acr acrosome, bv blood vessel, coe coelomic cavity, mes mesentery, nc nurse cell, nn nurse cell nucleus, nu sperm cell nucleus, Oo oocyte, on oocyte nucleus, sp spermatogonia, vnc ventral nerve cord" figureDoi="http://doi.org/10.5281/zenodo.12802038" httpUri="https://zenodo.org/record/12802038/files/figure.png" pageId="6" pageNumber="73">Fig. 5A, F</figureCitation>
). The complete reduction of the gut was observed in both juvenile (
<figureCitation id="135CBFD7FFE3FFFB6D88CA925CBBF939" box="[654,762,1720,1744]" captionStart="Fig" captionStartId="7.[136,165,1303,1324]" captionTargetBox="[144,1444,154,1255]" captionTargetId="figure-239@7.[144,1444,154,1255]" captionTargetPageId="7" captionText="Fig. 4 AZAN-stained paraffin histology. A Histological cross-section of a juvenile Aphrodita longipalpa featuring an endoparasitic immature Veneriserva pygoclava (denoted by a star). B Longitudinal section of V. pygoclava, highlighting the absence of a through gut, and continuous uninterrupted mesenteries. CH Cross-sections through the anterior region of V. pygoclava, showing the muscularized pharynx with jaws culminating in blind termination at section G. Abbreviations—ac acicula, br brain, df dorsal felt, el elytra, ja jaws, mo mouth, ne nephridium, pha pharynx, vnc ventral nerve cord" figureDoi="http://doi.org/10.5281/zenodo.12765015" httpUri="https://zenodo.org/record/12765015/files/figure.png" pageId="6" pageNumber="73">Fig. 4BH</figureCitation>
) and adult specimens of
<taxonomicName id="4C67D8D1FFE3FFFB6E89CAF05C5CF918" box="[399,541,1753,1777]" class="Polychaeta" family="Dorvilleidae" genus="Veneriserva" kingdom="Animalia" order="Eunicida" pageId="6" pageNumber="73" phylum="Annelida" rank="species" species="pygoclava">
<emphasis id="B9137F40FFE3FFFB6E89CAF05C5CF918" box="[399,541,1753,1777]" italics="true" pageId="6" pageNumber="73">V. pygoclava</emphasis>
</taxonomicName>
(
<figureCitation id="135CBFD7FFE3FFFB6D30CAF35CE1F918" box="[566,672,1753,1777]" captionStart="◂" captionStartId="10.[122,136,145,168]" captionTargetId="figure-13@9.[147,1440,154,1854]" captionTargetPageId="9" captionText="◂Fig.5 Gametogenesis in male and female Veneriserva pygoclava. AD Semi-thin histological sections of female Veneriserva pygoclava, stained with toluidine blue. A Cross-section of a female Veneriserva. B Close-up of large mature oocytes without discernible nurse cells. C Developing oocytes attached to mesenteries (mes), and oogonia proliferating from the ventral side of the dorsal blood vessel (bv). D Details of vitellogenic oocytes and nurse cells. Arrowheads indicate brownstained yolk platelets and yolk bodies. E Live sperm cells captured in a light micrograph. FG Cross-sections of male Veneriserva. Note the absence of a gut in the cross-sections. Abbreviations—ac acicula, acr acrosome, bv blood vessel, coe coelomic cavity, mes mesentery, nc nurse cell, nn nurse cell nucleus, nu sperm cell nucleus, Oo oocyte, on oocyte nucleus, sp spermatogonia, vnc ventral nerve cord" figureDoi="http://doi.org/10.5281/zenodo.12802038" httpUri="https://zenodo.org/record/12802038/files/figure.png" pageId="6" pageNumber="73">Fig. 5A, F</figureCitation>
) we sectioned histologically. Even though a jaw apparatus is present, and the animals can evert and move the maxillae in a pinching motion (
<figureCitation id="135CBFD7FFE3FFFB6E68CB175FFCF8BC" box="[366,445,1853,1877]" captionStart="◂" captionStartId="4.[122,136,145,168]" captionTargetId="figure-13@3.[147,1440,154,1854]" captionTargetPageId="3" captionText="◂Fig. 1 Live photos and dissection of parasitized Aphrodita longipalpa and Veneriserva pygoclava. A Ventral view of A. longipalpa. B Dorsal view of A. longipalpa with removed feltage chaetae, revealing the parasite visible through the body wall. C Ventrally dissected A. longipalpa, exposing the sizable female parasite. Veneriserva pygoclava individuals within the host are indicated by arrowheads. D Juvenile female V. pygoclava, with developing oocytes visible through the body wall along the mid-dorsal orange line. E Female V. pygoclava showing the mid-dorsal orange pigmentation and the white mark at the base of the prostomium. F Male V. pygoclava. G A large female and smaller male V. pygoclava, extracted from the same host. The pygidium is club-shaped in both males and females and juveniles. H Juvenile V. pygoclava shown from multiple angles, characterized by a complete white coloration; black jaws are magnified in panel" figureDoi="http://doi.org/10.5281/zenodo.12802034" httpUri="https://zenodo.org/record/12802034/files/figure.png" pageId="6" pageNumber="73">Fig. 1H</figureCitation>
). The maxillae and the fused mandibles appear reduced when compared to free-living
<taxonomicName id="4C67D8D1FFE3FFFB6C34CA5F5D86F964" authorityName="Chamberlin" authorityYear="1919" box="[818,967,1653,1677]" class="Polychaeta" family="Dorvilleidae" kingdom="Animalia" order="Eunicida" pageId="6" pageNumber="73" phylum="Annelida" rank="family">Dorvilleidae</taxonomicName>
. The pygidium of
<taxonomicName id="4C67D8D1FFE3FFFB6BA5CA5C5B78F964" box="[1187,1337,1653,1677]" class="Polychaeta" family="Dorvilleidae" genus="Veneriserva" kingdom="Animalia" order="Eunicida" pageId="6" pageNumber="73" phylum="Annelida" rank="species" species="pygoclava">
<emphasis id="B9137F40FFE3FFFB6BA5CA5C5B78F964" box="[1187,1337,1653,1677]" italics="true" pageId="6" pageNumber="73">V. pygoclava</emphasis>
</taxonomicName>
was confirmed as being club shaped, which inspired the species epithet. An anus, rudimentary or not, was not confirmed.
</paragraph>
</subSubSection>
<caption id="DF18F3DAFFE3FFFB6F8FC9BF5DD8F9FD" ID-DOI="http://doi.org/10.5281/zenodo.12765013" ID-Zenodo-Dep="12765013" httpUri="https://zenodo.org/record/12765013/files/figure.png" pageId="6" pageNumber="73" startId="6.[137,166,1429,1450]" subCaptionStartIDs="6.[1057,1184,1454,1476]" subCaptionStarts="Abbr" targetBox="[165,1423,154,1382]" targetPageId="6" targetType="figure">
<paragraph id="8BD8A352FFE3FFFB6F8FC9BF5DD8F9FD" blockId="6.[136,772,1428,1583]" lastBlockId="6.[816,1452,1427,1556]" pageId="6" pageNumber="73">
<emphasis id="B9137F40FFE3FFFB6F8FC9BF5EF9FA43" bold="true" box="[137,184,1429,1450]" pageId="6" pageNumber="73">Fig. 3</emphasis>
µCT visualization of parasites within
<taxonomicName id="4C67D8D1FFE3FFFB6D33C9BE5CBCFA40" authorityName="Essenberg" authorityYear="1917" box="[565,765,1428,1449]" class="Polychaeta" family="Aphroditidae" genus="Aphrodita" kingdom="Animalia" order="Phyllodocida" pageId="6" pageNumber="73" phylum="Annelida" rank="species" species="longipalpa">
<emphasis id="B9137F40FFE3FFFB6D33C9BE5CBCFA40" box="[565,765,1428,1449]" italics="true" pageId="6" pageNumber="73">Aphrodita longipalpa</emphasis>
</taxonomicName>
.
<emphasis id="B9137F40FFE3FFFB6F8EC9845ED9FA2A" bold="true" box="[136,152,1454,1475]" pageId="6" pageNumber="73">A</emphasis>
3D rendering of parasites shown within the projection of the host body.
<emphasis id="B9137F40FFE3FFFB6FC7C9E35EADFA37" bold="true" box="[193,236,1481,1502]" pageId="6" pageNumber="73">BD</emphasis>
Virtual dissections of surface renderings, showing crosssections of the host across three consecutive body regions, from anterior to posterior. Raw image data from the micro-CT stack, illustrating a horizontal section through the host (
<emphasis id="B9137F40FFE3FFFB6D3ECA335C06F9C7" bold="true" box="[568,583,1561,1582]" pageId="6" pageNumber="73">E</emphasis>
) and a sagittal section (
<emphasis id="B9137F40FFE3FFFB6C63C9BE5D32FA40" bold="true" box="[869,883,1428,1449]" pageId="6" pageNumber="73">F</emphasis>
). Head of the juvenile parasite is magnified to display the prominent jaws in white. Abbreviations—
<emphasis id="B9137F40FFE3FFFB6BB1C9855A88FA2D" box="[1207,1225,1455,1476]" italics="true" pageId="6" pageNumber="73">ja</emphasis>
jaws,
<emphasis id="B9137F40FFE3FFFB6A01C9855B5DFA2D" box="[1287,1308,1455,1476]" italics="true" pageId="6" pageNumber="73">ne</emphasis>
nephridia,
<emphasis id="B9137F40FFE3FFFB6A8FC9845BEAFA2A" box="[1417,1451,1454,1475]" italics="true" pageId="6" pageNumber="73">pha</emphasis>
pharynx. Female
<taxonomicName id="4C67D8D1FFE3FFFB6CD2C9E35AE4FA37" box="[980,1189,1481,1502]" class="Polychaeta" family="Dorvilleidae" genus="Veneriserva" kingdom="Animalia" order="Eunicida" pageId="6" pageNumber="73" phylum="Annelida" rank="species" species="pygoclava">
<emphasis id="B9137F40FFE3FFFB6CD2C9E35AE4FA37" box="[980,1189,1481,1502]" italics="true" pageId="6" pageNumber="73">Veneriserva pygoclava</emphasis>
</taxonomicName>
is shown in yellow or with yellow arrowheads and the juvenile
<taxonomicName id="4C67D8D1FFE3FFFB6B7AC9CE5AB1FA11" box="[1148,1264,1507,1529]" class="Polychaeta" family="Dorvilleidae" genus="Veneriserva" kingdom="Animalia" order="Eunicida" pageId="6" pageNumber="73" phylum="Annelida" rank="species" species="pygoclava">
<emphasis id="B9137F40FFE3FFFB6B7AC9CE5AB1FA11" box="[1148,1264,1507,1529]" italics="true" pageId="6" pageNumber="73">V. pygoclava</emphasis>
</taxonomicName>
in blue or with blue arrowheads
</paragraph>
</caption>
<subSubSection id="C37DF0D9FFE3FFFA6C36CAD35ABFF964" lastPageId="7" lastPageNumber="74" pageId="6" pageNumber="73" type="description">
<paragraph id="8BD8A352FFE3FFFB6C36CAD35A1CF8FD" blockId="6.[816,1117,1785,1812]" box="[816,1117,1785,1812]" pageId="6" pageNumber="73">
<heading id="D090143EFFE3FFFB6C36CAD35A1CF8FD" bold="true" box="[816,1117,1785,1812]" fontSize="11" level="1" pageId="6" pageNumber="73" reason="1">
<emphasis id="B9137F40FFE3FFFB6C36CAD35A1CF8FD" bold="true" box="[816,1117,1785,1812]" pageId="6" pageNumber="73">Epidermal ultrastructure</emphasis>
</heading>
</paragraph>
<paragraph id="8BD8A352FFE3FFFA6C36CB175ABFF964" blockId="6.[816,1451,1853,1911]" lastBlockId="7.[816,1453,1487,1677]" lastPageId="7" lastPageNumber="74" pageId="6" pageNumber="73">
The entire body surface of
<taxonomicName id="4C67D8D1FFE3FFFB6B4ECB145A8EF8BC" box="[1096,1231,1853,1877]" class="Polychaeta" family="Dorvilleidae" genus="Veneriserva" kingdom="Animalia" order="Eunicida" pageId="6" pageNumber="73" phylum="Annelida" rank="species" species="pygoclava">
<emphasis id="B9137F40FFE3FFFB6B4ECB145A8EF8BC" box="[1096,1231,1853,1877]" italics="true" pageId="6" pageNumber="73">V. pygoclava</emphasis>
</taxonomicName>
was densely covered with microvilli of the epidermal cells that pierced through the cuticle (
<figureCitation id="135CBFD7FFE2FFFA6E15C9E55FC2FA0E" box="[275,387,1487,1511]" captionStart="Fig" captionStartId="11.[136,165,1445,1466]" captionTargetBox="[144,1444,154,1398]" captionTargetId="figure-156@11.[144,1444,154,1398]" captionTargetPageId="11" captionText="Fig.6 Epidermal ultrastructure of Veneriserva pygoclava. AD TEM images of the epidermis revealing the presence of dense, modified microvilli (mv) that cover the body surface. A mucosecretory gland cell (gl) is discernable in A. D shows details of a multi-ciliated epidermal cell. B depicts the microvilli (mv) covering the cuticle (cu). Note the inflated tips of the microvilli and the electron-dense droplets. E Apically the epidermal cells display an abundance of transport vesicles (v). Arrowheads mark the branching microvilli piercing through the cuticle in all images. Abbreviations—ci cilia, m mitochondria, nc nucleus" figureDoi="http://doi.org/10.5281/zenodo.12765017" httpUri="https://zenodo.org/record/12765017/files/figure.png" pageId="7" pageNumber="74">Fig. 6AC</figureCitation>
). The cuticle was less than 500 µm in thickness. A thin electron-dense epicuticle was present and had an inner lighter and an outer denser zone. The outer electron dense zone of the epicuticle was covered with a thin layer of darkly stained particles, giving it a fuzzy appearance (
<figureCitation id="135CBFD7FFE2FFFA6FD7CA5F5F5CF964" box="[209,285,1653,1677]" captionStart="Fig" captionStartId="11.[136,165,1445,1466]" captionTargetBox="[144,1444,154,1398]" captionTargetId="figure-156@11.[144,1444,154,1398]" captionTargetPageId="11" captionText="Fig.6 Epidermal ultrastructure of Veneriserva pygoclava. AD TEM images of the epidermis revealing the presence of dense, modified microvilli (mv) that cover the body surface. A mucosecretory gland cell (gl) is discernable in A. D shows details of a multi-ciliated epidermal cell. B depicts the microvilli (mv) covering the cuticle (cu). Note the inflated tips of the microvilli and the electron-dense droplets. E Apically the epidermal cells display an abundance of transport vesicles (v). Arrowheads mark the branching microvilli piercing through the cuticle in all images. Abbreviations—ci cilia, m mitochondria, nc nucleus" figureDoi="http://doi.org/10.5281/zenodo.12765017" httpUri="https://zenodo.org/record/12765017/files/figure.png" pageId="7" pageNumber="74">Fig. 6B</figureCitation>
). The unbanded collagen fibers of the cuticle were embedded in an electron-light glycocalyx matrix and were more densely arranged towards the epicuticle (
<figureCitation id="135CBFD7FFE2FFFA6DA9CA925CB7F939" box="[687,758,1720,1744]" captionStart="Fig" captionStartId="11.[136,165,1445,1466]" captionTargetBox="[144,1444,154,1398]" captionTargetId="figure-156@11.[144,1444,154,1398]" captionTargetPageId="11" captionText="Fig.6 Epidermal ultrastructure of Veneriserva pygoclava. AD TEM images of the epidermis revealing the presence of dense, modified microvilli (mv) that cover the body surface. A mucosecretory gland cell (gl) is discernable in A. D shows details of a multi-ciliated epidermal cell. B depicts the microvilli (mv) covering the cuticle (cu). Note the inflated tips of the microvilli and the electron-dense droplets. E Apically the epidermal cells display an abundance of transport vesicles (v). Arrowheads mark the branching microvilli piercing through the cuticle in all images. Abbreviations—ci cilia, m mitochondria, nc nucleus" figureDoi="http://doi.org/10.5281/zenodo.12765017" httpUri="https://zenodo.org/record/12765017/files/figure.png" pageId="7" pageNumber="74">Fig. 6B</figureCitation>
). Distally, the epidermal cells contained numerous transport vesicles (inclusion bodies) (
<figureCitation id="135CBFD7FFE2FFFA6EBCCAD15C43F8FA" box="[442,514,1787,1811]" captionStart="Fig" captionStartId="11.[136,165,1445,1466]" captionTargetBox="[144,1444,154,1398]" captionTargetId="figure-156@11.[144,1444,154,1398]" captionTargetPageId="11" captionText="Fig.6 Epidermal ultrastructure of Veneriserva pygoclava. AD TEM images of the epidermis revealing the presence of dense, modified microvilli (mv) that cover the body surface. A mucosecretory gland cell (gl) is discernable in A. D shows details of a multi-ciliated epidermal cell. B depicts the microvilli (mv) covering the cuticle (cu). Note the inflated tips of the microvilli and the electron-dense droplets. E Apically the epidermal cells display an abundance of transport vesicles (v). Arrowheads mark the branching microvilli piercing through the cuticle in all images. Abbreviations—ci cilia, m mitochondria, nc nucleus" figureDoi="http://doi.org/10.5281/zenodo.12765017" httpUri="https://zenodo.org/record/12765017/files/figure.png" pageId="7" pageNumber="74">Fig. 6E</figureCitation>
). Individual mucosecretory cells were abundant in the epidermis and were situated between supportive cells (
<figureCitation id="135CBFD7FFE2FFFA6EB1CB175C48F8BC" box="[439,521,1853,1877]" captionStart="Fig" captionStartId="11.[136,165,1445,1466]" captionTargetBox="[144,1444,154,1398]" captionTargetId="figure-156@11.[144,1444,154,1398]" captionTargetPageId="11" captionText="Fig.6 Epidermal ultrastructure of Veneriserva pygoclava. AD TEM images of the epidermis revealing the presence of dense, modified microvilli (mv) that cover the body surface. A mucosecretory gland cell (gl) is discernable in A. D shows details of a multi-ciliated epidermal cell. B depicts the microvilli (mv) covering the cuticle (cu). Note the inflated tips of the microvilli and the electron-dense droplets. E Apically the epidermal cells display an abundance of transport vesicles (v). Arrowheads mark the branching microvilli piercing through the cuticle in all images. Abbreviations—ci cilia, m mitochondria, nc nucleus" figureDoi="http://doi.org/10.5281/zenodo.12765017" httpUri="https://zenodo.org/record/12765017/files/figure.png" pageId="7" pageNumber="74">Fig. 6A</figureCitation>
). Patches of multiciliated epidermal cells were present scattered around the body (
<figureCitation id="135CBFD7FFE2FFFA6C47C9E55DCDFA0E" box="[833,908,1487,1511]" captionStart="Fig" captionStartId="11.[136,165,1445,1466]" captionTargetBox="[144,1444,154,1398]" captionTargetId="figure-156@11.[144,1444,154,1398]" captionTargetPageId="11" captionText="Fig.6 Epidermal ultrastructure of Veneriserva pygoclava. AD TEM images of the epidermis revealing the presence of dense, modified microvilli (mv) that cover the body surface. A mucosecretory gland cell (gl) is discernable in A. D shows details of a multi-ciliated epidermal cell. B depicts the microvilli (mv) covering the cuticle (cu). Note the inflated tips of the microvilli and the electron-dense droplets. E Apically the epidermal cells display an abundance of transport vesicles (v). Arrowheads mark the branching microvilli piercing through the cuticle in all images. Abbreviations—ci cilia, m mitochondria, nc nucleus" figureDoi="http://doi.org/10.5281/zenodo.12765017" httpUri="https://zenodo.org/record/12765017/files/figure.png" pageId="7" pageNumber="74">Fig. 6D</figureCitation>
). These multiciliated cells contained larger amounts of mitochondria and were also covered with microvilli (
<figureCitation id="135CBFD7FFE2FFFA6C44CA3B5DD3F9C0" box="[834,914,1553,1577]" captionStart="Fig" captionStartId="11.[136,165,1445,1466]" captionTargetBox="[144,1444,154,1398]" captionTargetId="figure-156@11.[144,1444,154,1398]" captionTargetPageId="11" captionText="Fig.6 Epidermal ultrastructure of Veneriserva pygoclava. AD TEM images of the epidermis revealing the presence of dense, modified microvilli (mv) that cover the body surface. A mucosecretory gland cell (gl) is discernable in A. D shows details of a multi-ciliated epidermal cell. B depicts the microvilli (mv) covering the cuticle (cu). Note the inflated tips of the microvilli and the electron-dense droplets. E Apically the epidermal cells display an abundance of transport vesicles (v). Arrowheads mark the branching microvilli piercing through the cuticle in all images. Abbreviations—ci cilia, m mitochondria, nc nucleus" figureDoi="http://doi.org/10.5281/zenodo.12765017" httpUri="https://zenodo.org/record/12765017/files/figure.png" pageId="7" pageNumber="74">Fig. 6D</figureCitation>
). Microvilli were branched (
<figureCitation id="135CBFD7FFE2FFFA6BDECA3B5B62F9C0" box="[1240,1315,1553,1577]" captionStart="Fig" captionStartId="11.[136,165,1445,1466]" captionTargetBox="[144,1444,154,1398]" captionTargetId="figure-156@11.[144,1444,154,1398]" captionTargetPageId="11" captionText="Fig.6 Epidermal ultrastructure of Veneriserva pygoclava. AD TEM images of the epidermis revealing the presence of dense, modified microvilli (mv) that cover the body surface. A mucosecretory gland cell (gl) is discernable in A. D shows details of a multi-ciliated epidermal cell. B depicts the microvilli (mv) covering the cuticle (cu). Note the inflated tips of the microvilli and the electron-dense droplets. E Apically the epidermal cells display an abundance of transport vesicles (v). Arrowheads mark the branching microvilli piercing through the cuticle in all images. Abbreviations—ci cilia, m mitochondria, nc nucleus" figureDoi="http://doi.org/10.5281/zenodo.12765017" httpUri="https://zenodo.org/record/12765017/files/figure.png" pageId="7" pageNumber="74">Fig. 6E</figureCitation>
) and had an enlarged, inflated tip covered with electron-dense droplets (
<figureCitation id="135CBFD7FFE2FFFA6C44CA7E5DCAF985" box="[834,907,1620,1644]" captionStart="Fig" captionStartId="11.[136,165,1445,1466]" captionTargetBox="[144,1444,154,1398]" captionTargetId="figure-156@11.[144,1444,154,1398]" captionTargetPageId="11" captionText="Fig.6 Epidermal ultrastructure of Veneriserva pygoclava. AD TEM images of the epidermis revealing the presence of dense, modified microvilli (mv) that cover the body surface. A mucosecretory gland cell (gl) is discernable in A. D shows details of a multi-ciliated epidermal cell. B depicts the microvilli (mv) covering the cuticle (cu). Note the inflated tips of the microvilli and the electron-dense droplets. E Apically the epidermal cells display an abundance of transport vesicles (v). Arrowheads mark the branching microvilli piercing through the cuticle in all images. Abbreviations—ci cilia, m mitochondria, nc nucleus" figureDoi="http://doi.org/10.5281/zenodo.12765017" httpUri="https://zenodo.org/record/12765017/files/figure.png" pageId="7" pageNumber="74">Fig. 6B</figureCitation>
). The inflated portion of a microvillus was bulbous and had a maximum diameter of ±
<quantity id="4C9F0EB7FFE2FFFA6BA3CA5F5ABBF964" box="[1189,1274,1653,1677]" metricMagnitude="-7" metricUnit="m" metricValue="3.0" pageId="7" pageNumber="74" unit="nm" value="300.0">300 nm</quantity>
.
</paragraph>
</subSubSection>
<caption id="DF18F3DAFFE2FFFA6F8EC93D5B0DFA92" ID-DOI="http://doi.org/10.5281/zenodo.12765015" ID-Zenodo-Dep="12765015" httpUri="https://zenodo.org/record/12765015/files/figure.png" pageId="7" pageNumber="74" startId="7.[136,165,1303,1324]" subCaptionStartIDs="7.[1388,1452,1328,1350]" subCaptionStarts="Abbr" targetBox="[144,1444,154,1255]" targetPageId="7" targetType="figure">
<paragraph id="8BD8A352FFE2FFFA6F8EC93D5B0DFA92" blockId="7.[136,771,1301,1430]" lastBlockId="7.[816,1452,1301,1403]" pageId="7" pageNumber="74">
<emphasis id="B9137F40FFE2FFFA6F8EC93D5EF9FAC5" bold="true" box="[136,184,1303,1324]" pageId="7" pageNumber="74">Fig. 4</emphasis>
AZAN-stained paraffin histology.
<emphasis id="B9137F40FFE2FFFA6D07C93F5C50FAC3" bold="true" box="[513,529,1301,1322]" pageId="7" pageNumber="74">A</emphasis>
Histological cross-section of a juvenile
<taxonomicName id="4C67D8D1FFE2FFFA6E02C91A5F88FAAC" authorityName="Essenberg" authorityYear="1917" box="[260,457,1328,1349]" class="Polychaeta" family="Aphroditidae" genus="Aphrodita" kingdom="Animalia" order="Phyllodocida" pageId="7" pageNumber="74" phylum="Annelida" rank="species" species="longipalpa">
<emphasis id="B9137F40FFE2FFFA6E02C91A5F88FAAC" box="[260,457,1328,1349]" italics="true" pageId="7" pageNumber="74">Aphrodita longipalpa</emphasis>
</taxonomicName>
featuring an endoparasitic immature
<taxonomicName id="4C67D8D1FFE2FFFA6FB5C9615FC5FA89" box="[179,388,1355,1376]" class="Polychaeta" family="Dorvilleidae" genus="Veneriserva" kingdom="Animalia" order="Eunicida" pageId="7" pageNumber="74" phylum="Annelida" rank="species" species="pygoclava">
<emphasis id="B9137F40FFE2FFFA6FB5C9615FC5FA89" box="[179,388,1355,1376]" italics="true" pageId="7" pageNumber="74">Veneriserva pygoclava</emphasis>
</taxonomicName>
(denoted by a star).
<emphasis id="B9137F40FFE2FFFA6D4CC9615C18FA89" bold="true" box="[586,601,1355,1376]" pageId="7" pageNumber="74">B</emphasis>
Longitudinal section of
<taxonomicName id="4C67D8D1FFE2FFFA6FC8C94C5F05FA93" box="[206,324,1381,1403]" class="Polychaeta" family="Dorvilleidae" genus="Veneriserva" kingdom="Animalia" order="Eunicida" pageId="7" pageNumber="74" phylum="Annelida" rank="species" species="pygoclava">
<emphasis id="B9137F40FFE2FFFA6FC8C94C5F05FA93" box="[206,324,1381,1403]" italics="true" pageId="7" pageNumber="74">V. pygoclava</emphasis>
</taxonomicName>
, highlighting the absence of a through gut, and continuous uninterrupted mesenteries.
<emphasis id="B9137F40FFE2FFFA6EF1C9AA5C65FA7F" bold="true" box="[503,548,1408,1430]" pageId="7" pageNumber="74">CH</emphasis>
Cross-sections through the anterior region of
<taxonomicName id="4C67D8D1FFE2FFFA6CF9C93C5A34FAC3" box="[1023,1141,1301,1323]" class="Polychaeta" family="Dorvilleidae" genus="Veneriserva" kingdom="Animalia" order="Eunicida" pageId="7" pageNumber="74" phylum="Annelida" rank="species" species="pygoclava">
<emphasis id="B9137F40FFE2FFFA6CF9C93C5A34FAC3" box="[1023,1141,1301,1323]" italics="true" pageId="7" pageNumber="74">V. pygoclava</emphasis>
</taxonomicName>
, showing the muscularized pharynx with jaws culminating in blind termination at section
<emphasis id="B9137F40FFE2FFFA6A49C91A5B1EFAAF" bold="true" box="[1359,1375,1328,1350]" pageId="7" pageNumber="74">G</emphasis>
. Abbreviations—
<emphasis id="B9137F40FFE2FFFA6C89C9615DE5FA89" box="[911,932,1355,1376]" italics="true" pageId="7" pageNumber="74">ac</emphasis>
acicula,
<emphasis id="B9137F40FFE2FFFA6CFBC9615A50FA89" box="[1021,1041,1355,1376]" italics="true" pageId="7" pageNumber="74">br</emphasis>
brain,
<emphasis id="B9137F40FFE2FFFA6B5EC9615A2BFA89" box="[1112,1130,1355,1376]" italics="true" pageId="7" pageNumber="74">df</emphasis>
dorsal felt,
<emphasis id="B9137F40FFE2FFFA6BE7C9615AB0FA89" box="[1249,1265,1355,1376]" italics="true" pageId="7" pageNumber="74">el</emphasis>
elytra,
<emphasis id="B9137F40FFE2FFFA6A3BC9615B0EFA89" box="[1341,1359,1355,1376]" italics="true" pageId="7" pageNumber="74">ja</emphasis>
jaws,
<emphasis id="B9137F40FFE2FFFA6A96C9615BEDFA89" box="[1424,1452,1355,1376]" italics="true" pageId="7" pageNumber="74">mo</emphasis>
mouth,
<emphasis id="B9137F40FFE2FFFA6C70C94C5DCAFA92" box="[886,907,1382,1403]" italics="true" pageId="7" pageNumber="74">ne</emphasis>
nephridium,
<emphasis id="B9137F40FFE2FFFA6B03C94F5A66FA93" box="[1029,1063,1381,1402]" italics="true" pageId="7" pageNumber="74">pha</emphasis>
pharynx,
<emphasis id="B9137F40FFE2FFFA6B85C94C5AE3FA92" box="[1155,1186,1382,1403]" italics="true" pageId="7" pageNumber="74">vnc</emphasis>
ventral nerve cord
</paragraph>
</caption>
<subSubSection id="C37DF0D9FFE2FFF56C36CA9C5CAFFD2E" lastPageId="8" lastPageNumber="75" pageId="7" pageNumber="74" type="discussion">
<paragraph id="8BD8A352FFE2FFFA6C36CA9C5B46F938" blockId="7.[816,1287,1718,1745]" box="[816,1287,1718,1745]" pageId="7" pageNumber="74">
<heading id="D090143EFFE2FFFA6C36CA9C5B46F938" bold="true" box="[816,1287,1718,1745]" fontSize="11" level="1" pageId="7" pageNumber="74" reason="1">
<emphasis id="B9137F40FFE2FFFA6C36CA9C5B46F938" bold="true" box="[816,1287,1718,1745]" pageId="7" pageNumber="74">
Phylogenetic placement of
<taxonomicName id="4C67D8D1FFE2FFFA6B7DCA9C5B46F938" box="[1147,1287,1718,1745]" class="Polychaeta" family="Dorvilleidae" genus="Veneriserva" kingdom="Animalia" order="Eunicida" pageId="7" pageNumber="74" phylum="Annelida" rank="genus">
<emphasis id="B9137F40FFE2FFFA6B7DCA9C5B46F938" bold="true" box="[1147,1287,1718,1745]" italics="true" pageId="7" pageNumber="74">Veneriserva</emphasis>
</taxonomicName>
</emphasis>
</heading>
</paragraph>
<paragraph id="8BD8A352FFE2FFF56C36CAD15CAFFD2E" blockId="7.[816,1452,1787,1911]" lastBlockId="8.[135,772,153,711]" lastPageId="8" lastPageNumber="75" pageId="7" pageNumber="74">
The ML analysis of the three mitochondrial and two nuclear genes for
<taxonomicName id="4C67D8D1FFE2FFFA6CA6CB365A6AF8DD" authorityName="Chamberlin" authorityYear="1919" box="[928,1067,1820,1844]" class="Polychaeta" family="Dorvilleidae" kingdom="Animalia" order="Eunicida" pageId="7" pageNumber="74" phylum="Annelida" rank="family">Dorvilleidae</taxonomicName>
showed two main clades (labeled A and B) containing all
<taxonomicName id="4C67D8D1FFE2FFFA6B4ECB175AA5F8BC" authorityName="Claparede &amp; Mecznikow" authorityYear="1869" box="[1096,1252,1853,1877]" class="Polychaeta" family="Dorvilleidae" genus="Ophryotrocha" kingdom="Animalia" order="Eunicida" pageId="7" pageNumber="74" phylum="Annelida" rank="genus">
<emphasis id="B9137F40FFE2FFFA6B4ECB175AA5F8BC" box="[1096,1252,1853,1877]" italics="true" pageId="7" pageNumber="74">Ophryotrocha</emphasis>
</taxonomicName>
terminals as well as members of other genera such as
<taxonomicName id="4C67D8D1FFE2FFFA6BCDCB755B7BF89E" authorityName="Jumars" authorityYear="1974" box="[1227,1338,1887,1911]" class="Polychaeta" family="Dorvilleidae" genus="Exallopus" kingdom="Animalia" order="Eunicida" pageId="7" pageNumber="74" phylum="Annelida" rank="genus">
<emphasis id="B9137F40FFE2FFFA6BCDCB755B7BF89E" box="[1227,1338,1887,1911]" italics="true" pageId="7" pageNumber="74">Exallopus</emphasis>
</taxonomicName>
,
<taxonomicName id="4C67D8D1FFE2FFFA6A4ECB755BE5F89E" authorityName="Marenzeller" authorityYear="1902" box="[1352,1444,1887,1911]" class="Polychaeta" family="Dorvilleidae" genus="Iphitime" kingdom="Animalia" order="Eunicida" pageId="7" pageNumber="74" phylum="Annelida" rank="genus">
<emphasis id="B9137F40FFE2FFFA6A4ECB755BE5F89E" box="[1352,1444,1887,1911]" italics="true" pageId="7" pageNumber="74">Iphitime</emphasis>
</taxonomicName>
,
<taxonomicName id="4C67D8D1FFEDFFF56F8ECCB35F20FF58" authorityName="Hilbig &amp; Blake" authorityYear="1991" box="[136,353,153,177]" class="Polychaeta" family="Dorvilleidae" genus="Pseudophryotrocha" kingdom="Animalia" order="Eunicida" pageId="8" pageNumber="75" phylum="Annelida" rank="genus">
<emphasis id="B9137F40FFEDFFF56F8ECCB35F20FF58" box="[136,353,153,177]" italics="true" pageId="8" pageNumber="75">Pseudophryotrocha</emphasis>
</taxonomicName>
, and
<taxonomicName id="4C67D8D1FFEDFFF56EA2CCB05C68FF58" box="[420,553,154,177]" class="Polychaeta" family="Dorvilleidae" genus="Veneriserva" kingdom="Animalia" order="Eunicida" pageId="8" pageNumber="75" phylum="Annelida" rank="genus">
<emphasis id="B9137F40FFEDFFF56EA2CCB05C68FF58" box="[420,553,154,177]" italics="true" pageId="8" pageNumber="75">Veneriserva</emphasis>
</taxonomicName>
.
<taxonomicName id="4C67D8D1FFEDFFF56D31CCB05E85FF3A" class="Polychaeta" family="Dorvilleidae" genus="Veneriserva" kingdom="Animalia" order="Eunicida" pageId="8" pageNumber="75" phylum="Annelida" rank="species" species="pygoclava">
<emphasis id="B9137F40FFEDFFF56D31CCB05E85FF3A" italics="true" pageId="8" pageNumber="75">Veneriserva pygoclava</emphasis>
</taxonomicName>
was in clade B with a relatively long branch and formed a poorly supported clade with the three included terminals of
<taxonomicName id="4C67D8D1FFEDFFF56E17CCD75F2AFEFC" authorityName="Marenzeller" authorityYear="1902" box="[273,363,253,277]" class="Polychaeta" family="Dorvilleidae" genus="Iphitime" kingdom="Animalia" order="Eunicida" pageId="8" pageNumber="75" phylum="Annelida" rank="genus">
<emphasis id="B9137F40FFEDFFF56E17CCD75F2AFEFC" box="[273,363,253,277]" italics="true" pageId="8" pageNumber="75">Iphitime</emphasis>
</taxonomicName>
(
<figureCitation id="135CBFD7FFEDFFF56E85CCD75FF8FEFC" box="[387,441,253,277]" captionStart="Fig" captionStartId="12.[138,167,1427,1448]" captionTargetBox="[144,1444,156,1379]" captionTargetId="figure-210@12.[143,1444,154,1380]" captionTargetPageId="12" captionText="Fig.7 Maximum likelihood (ML) tree of Dorvilleidae. The tree depicts the phylogenetic relationships within Dorvilleidae inferred through concatenated 16S, COI, Cytb, 18S, and H3 sequences. Bootstrap support values are provided for each node. Nodes with complete support are indicated with an asterisk (*); values below 50% are not shown. Branches of parasitic/symbiotic species highlighted in blue. Haplotype network for 5 Veneriserva pygoclava specimens is shown next to the tree" figureDoi="http://doi.org/10.5281/zenodo.12765019" httpUri="https://zenodo.org/record/12765019/files/figure.png" pageId="8" pageNumber="75">Fig. 7</figureCitation>
), which are all symbiotic with Crustacea (
<bibRefCitation id="EFF6DEA3FFEDFFF56E0CCD355C69FEDE" author="de Paiva, P. C. &amp; Nonato, E. F." box="[266,552,287,311]" pageId="8" pageNumber="75" pagination="209 - 215" refId="ref9199" refString="de Paiva, P. C., &amp; Nonato, E. F. % 1991). On the genus Iphitime % Polychaeta: Iphitimidae) and description of Iphitime sartorae sp. nov. a commensal of brachyuran crabs. Ophelia, 34 % 3), 209 - 215. https: // doi. org / 10.1080 / 00785326.1991. 10429696" type="journal article" year="1991">de Paiva and Nonato, 1991</bibRefCitation>
). It was sister taxon to
<taxonomicName id="4C67D8D1FFEDFFF56FA0CD6A5C22FEB1" authority="Fage &amp; Legendre, 1934" authorityName="Fage &amp; Legendre" authorityYear="1934" box="[166,611,320,345]" class="Polychaeta" family="Dorvilleidae" genus="Iphitime" kingdom="Animalia" order="Eunicida" pageId="8" pageNumber="75" phylum="Annelida" rank="species" species="paguri">
<emphasis id="B9137F40FFEDFFF56FA0CD6A5F12FEB1" box="[166,339,320,344]" italics="true" pageId="8" pageNumber="75">Iphitime paguri</emphasis>
<bibRefCitation id="EFF6DEA3FFEDFFF56E5ACD6B5C22FEB1" author="Fage, L. &amp; Legendre, R." box="[348,611,320,345]" pageId="8" pageNumber="75" pagination="299 - 305" refId="ref9645" refString="Fage, L., &amp; Legendre, R. % 1934). Les annelides polychetes du genre Iphitime. A propos d'une espece nouvelle commensale des pagures, Iphitime paguri n. sp. Bulletin De La Societe Zoologique De France, 58, 299 - 305." type="journal article" year="1934">Fage &amp; Legendre, 1934</bibRefCitation>
</taxonomicName>
, making
<taxonomicName id="4C67D8D1FFEDFFF56DC9CD6A5EF6FE90" authorityName="Marenzeller" authorityYear="1902" class="Polychaeta" family="Dorvilleidae" genus="Iphitime" kingdom="Animalia" order="Eunicida" pageId="8" pageNumber="75" phylum="Annelida" rank="genus">
<emphasis id="B9137F40FFEDFFF56DC9CD6A5EF6FE90" italics="true" pageId="8" pageNumber="75">Iphitime</emphasis>
</taxonomicName>
paraphyletic, though there was little support for this. This
<taxonomicName id="4C67D8D1FFEDFFF56FC7CDA95F5CFE72" authorityName="Marenzeller" authorityYear="1902" box="[193,285,387,411]" class="Polychaeta" family="Dorvilleidae" genus="Iphitime" kingdom="Animalia" order="Eunicida" pageId="8" pageNumber="75" phylum="Annelida" rank="genus">
<emphasis id="B9137F40FFEDFFF56FC7CDA95F5CFE72" box="[193,285,387,411]" italics="true" pageId="8" pageNumber="75">Iphitime</emphasis>
</taxonomicName>
/
<taxonomicName id="4C67D8D1FFEDFFF56E23CDAE5FE8FE72" box="[293,425,388,411]" class="Polychaeta" family="Dorvilleidae" genus="Veneriserva" kingdom="Animalia" order="Eunicida" pageId="8" pageNumber="75" phylum="Annelida" rank="genus">
<emphasis id="B9137F40FFEDFFF56E23CDAE5FE8FE72" box="[293,425,388,411]" italics="true" pageId="8" pageNumber="75">Veneriserva</emphasis>
</taxonomicName>
clade formed a well-supported clade together with a sister clade of
<taxonomicName id="4C67D8D1FFEDFFF56D1ECD8E5CF0FE55" authorityName="Claparede &amp; Mecznikow" authorityYear="1869" box="[536,689,420,444]" class="Polychaeta" family="Dorvilleidae" genus="Ophryotrocha" kingdom="Animalia" order="Eunicida" pageId="8" pageNumber="75" phylum="Annelida" rank="genus">
<emphasis id="B9137F40FFEDFFF56D1ECD8E5CF0FE55" box="[536,689,420,444]" italics="true" pageId="8" pageNumber="75">Ophryotrocha</emphasis>
</taxonomicName>
mainly associated with hydrothermal vents (6 species) (
<bibRefCitation id="EFF6DEA3FFEDFFF56D8CCDEF5EFFFE16" author="Zhang, D. &amp; Zhou, Y. &amp; Yen, N. &amp; Hiley, A. S. &amp; Rouse, G. W." pageId="8" pageNumber="75" pagination="167 - 194" refId="ref12523" refString="Zhang, D., Zhou, Y., Yen, N., Hiley, A. S., &amp; Rouse, G. W. % 2023). Ophryotrocha % Dorvilleidae, Polychaeta, Annelida) from deep-sea hydrothermal vents, with the description of five new species. European Journal of Taxonomy, 864, 167 - 194. https: // doi. org / 10. 5852 / ejt. 2023.864. 2101" type="journal article" year="2023">Zhang et al., 2023</bibRefCitation>
) and one with a whale fall
<taxonomicName id="4C67D8D1FFEDFFF56EFFCDCD5EFEFDC9" authority="Taboada et al., 2013" authorityName="Taboada" authorityYear="2013" class="Polychaeta" family="Dorvilleidae" genus="Ophryotrocha" kingdom="Animalia" order="Eunicida" pageId="8" pageNumber="75" phylum="Annelida" rank="species" species="clava">
<emphasis id="B9137F40FFEDFFF56EFFCDCD5C16FE16" box="[505,599,487,511]" italics="true" pageId="8" pageNumber="75">O. clava</emphasis>
<bibRefCitation id="EFF6DEA3FFEDFFF56D59CDCD5EFEFDC9" author="Taboada, S. &amp; Wiklund, H. &amp; Glover, A. G. &amp; Dahlgren, T. G. &amp; Cristobo, J. &amp; Avila, C." pageId="8" pageNumber="75" pagination="1031 - 1045" refId="ref11760" refString="Taboada, S., Wiklund, H., Glover, A. G., Dahlgren, T. G., Cristobo, J., &amp; Avila, C. % 2013). Two new Antarctic Ophryotrocha % Annelida: Dorvilleidae) described from shallow-water whale bones. Polar Biology, 36, 1031 - 1045. https: // doi. org / 10.1007 / s 00300 - 013 - 1326 - 4" type="journal article" year="2013">Taboada et al., 2013</bibRefCitation>
</taxonomicName>
. Another clade of symbiotic dorvilleids, also living with Crustacea (
<taxonomicName id="4C67D8D1FFEDFFF56E3FCE005CF7FDAB" authority="Martin et al., 1991" authorityName="Martin" authorityYear="1991" box="[313,694,553,578]" class="Polychaeta" family="Dorvilleidae" genus="Ophryotrocha" kingdom="Animalia" order="Eunicida" pageId="8" pageNumber="75" phylum="Annelida" rank="species" species="mediterranea">
<emphasis id="B9137F40FFEDFFF56E3FCE005FA8FDA8" box="[313,489,553,577]" italics="true" pageId="8" pageNumber="75">O. mediterranea</emphasis>
<bibRefCitation id="EFF6DEA3FFEDFFF56EE9CE035CF7FDAB" author="Martin, D. &amp; Abello, P. &amp; Cartes, J." box="[495,694,553,578]" pageId="8" pageNumber="75" pagination="279 - 292" refId="ref10604" refString="Martin, D., Abello, P., &amp; Cartes, J. % 1991). A new species of Ophryotrocha % Polychaeta: Dorvilleidae) commensal in Geryon longipes % Crustacea: Brachyura) from the western Mediterranean Sea. Journal of Natural History, 25 % 2), 279 - 292. https: // doi. org / 10. 1080 / 00222939100770201" type="journal article" year="1991">Martin et al., 1991</bibRefCitation>
</taxonomicName>
and
<taxonomicName id="4C67D8D1FFEDFFF56DEFCE005FEAFD8A" authority="Esmark, 1874" authorityName="Esmark" authorityYear="1874" class="Polychaeta" family="Dorvilleidae" genus="Ophryotrocha" kingdom="Animalia" order="Eunicida" pageId="8" pageNumber="75" phylum="Annelida" rank="species" species="geryonicola">
<emphasis id="B9137F40FFEDFFF56DEFCE005F4BFD8A" italics="true" pageId="8" pageNumber="75">O. geryonicola</emphasis>
<bibRefCitation id="EFF6DEA3FFEDFFF56E14CE615FEAFD8A" author="Esmark, L." box="[274,427,587,611]" pageId="8" pageNumber="75" pagination="497 - 498" refId="ref9591" refString="Esmark, L. % 1874). Forhandlinger fra Videnskabs-Selskabet i Christiania. 497 - 498." type="book chapter" year="1874">Esmark, 1874</bibRefCitation>
</taxonomicName>
), formed a clade with the free-living
<taxonomicName id="4C67D8D1FFEDFFF56FCACE475F8AFD6D" authority="Banse, 1963" authorityName="Banse" authorityYear="1963" box="[204,459,620,644]" class="Polychaeta" family="Dorvilleidae" genus="Ophryotrocha" kingdom="Animalia" order="Eunicida" pageId="8" pageNumber="75" phylum="Annelida" rank="species" species="vivipara">
<emphasis id="B9137F40FFEDFFF56FCACE475F02FD6D" box="[204,323,621,644]" italics="true" pageId="8" pageNumber="75">O. vivipara</emphasis>
<bibRefCitation id="EFF6DEA3FFEDFFF56E4FCE475F8AFD6D" author="Banse, K." box="[329,459,620,644]" pageId="8" pageNumber="75" pagination="197 - 208" refId="ref8665" refString="Banse, K. % 1963). Polychaetous annelids from Puget Sound and the San Juan Archipelago, Washington. Proceedings of the Biological Society of Washington, 76, 197 - 208." type="journal article" year="1963">Banse, 1963</bibRefCitation>
</taxonomicName>
(
<figureCitation id="135CBFD7FFEDFFF56EE4CE465C56FD6D" box="[482,535,620,644]" captionStart="Fig" captionStartId="12.[138,167,1427,1448]" captionTargetBox="[144,1444,156,1379]" captionTargetId="figure-210@12.[143,1444,154,1380]" captionTargetPageId="12" captionText="Fig.7 Maximum likelihood (ML) tree of Dorvilleidae. The tree depicts the phylogenetic relationships within Dorvilleidae inferred through concatenated 16S, COI, Cytb, 18S, and H3 sequences. Bootstrap support values are provided for each node. Nodes with complete support are indicated with an asterisk (*); values below 50% are not shown. Branches of parasitic/symbiotic species highlighted in blue. Haplotype network for 5 Veneriserva pygoclava specimens is shown next to the tree" figureDoi="http://doi.org/10.5281/zenodo.12765019" httpUri="https://zenodo.org/record/12765019/files/figure.png" pageId="8" pageNumber="75">Fig. 7</figureCitation>
) and was also in clade B.
<specimenCount id="9D6168DBFFEDFFF56FA0CEA75F0AFD4C" box="[166,331,653,677]" count="5" pageId="8" pageNumber="75" type="generic">Five specimens</specimenCount>
of
<taxonomicName id="4C67D8D1FFEDFFF56E69CEA45FB9FD4C" box="[367,504,653,677]" class="Polychaeta" family="Dorvilleidae" genus="Veneriserva" kingdom="Animalia" order="Eunicida" pageId="8" pageNumber="75" phylum="Annelida" rank="species" species="pygoclava">
<emphasis id="B9137F40FFEDFFF56E69CEA45FB9FD4C" box="[367,504,653,677]" italics="true" pageId="8" pageNumber="75">V. pygoclava</emphasis>
</taxonomicName>
exhibited four CO1 haplotypes, each differing by a few mutational steps (
<figureCitation id="135CBFD7FFEDFFF56DAECE855C9FFD2F" box="[680,734,687,711]" captionStart="Fig" captionStartId="12.[138,167,1427,1448]" captionTargetBox="[144,1444,156,1379]" captionTargetId="figure-210@12.[143,1444,154,1380]" captionTargetPageId="12" captionText="Fig.7 Maximum likelihood (ML) tree of Dorvilleidae. The tree depicts the phylogenetic relationships within Dorvilleidae inferred through concatenated 16S, COI, Cytb, 18S, and H3 sequences. Bootstrap support values are provided for each node. Nodes with complete support are indicated with an asterisk (*); values below 50% are not shown. Branches of parasitic/symbiotic species highlighted in blue. Haplotype network for 5 Veneriserva pygoclava specimens is shown next to the tree" figureDoi="http://doi.org/10.5281/zenodo.12765019" httpUri="https://zenodo.org/record/12765019/files/figure.png" pageId="8" pageNumber="75">Fig. 7</figureCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>