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<document id="DFDB6ECF2CCA81D0AD2992DDFC644CFC" ID-DOI="10.1080/0022293031000155395" ID-ISSN="1464-5262" ID-Zenodo-Dep="5250248" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.taxonomicNames_approvedBy="felipe" checkinTime="1629849994237" checkinUser="felipe" docAuthor="Williams, Jason" docDate="2004" docId="537A7F7AFF94FD2B564E9B71707DFBE9" docLanguage="en" docName="JNatHist.38.11.1339-1358.pdf" docOrigin="Journal of Natural History 38 (11)" docSource="http://www.informaworld.com/openurl?genre=article&amp;doi=10.1080/0022293031000155395&amp;magic=crossref||D404A21C5BB053405B1A640AFFD44AE3" docStyle="DocumentStyle:443144A73DAE3E092C51A378E4CD0138.2:JNatHist.2003-2004.journal_article.1cover" docStyleId="443144A73DAE3E092C51A378E4CD0138" docStyleName="JNatHist.2003-2004.journal_article.1cover" docStyleVersion="2" docTitle="Polydorella kamakamai Williams 2004, sp. n." docType="treatment" docVersion="3" lastPageNumber="1348" masterDocId="AF430702FF90FD21544F9F66742AFFD1" masterDocTitle="Reproduction and morphology of Polydorella (Polychaeta: Spionidae), including the description of a new species from the Philippines" masterLastPageNumber="1358" masterPageNumber="1339" pageNumber="1342" updateTime="1699120712116" updateUser="plazi" zenodo-license-document="CLOSED">
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<mods:title id="DA49323DA1300B6D24F8D9A863696989">Reproduction and morphology of Polydorella (Polychaeta: Spionidae), including the description of a new species from the Philippines</mods:title>
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<mods:namePart id="8AF4CB3DBAF027C64157B2A20FAC93C0">Williams, Jason</mods:namePart>
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<mods:title id="1B2A43EFDB3106026DF4C8570A160E04">Journal of Natural History</mods:title>
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<mods:date id="7FB843374C3DC3631AD53FBF9F07F4E5">2004</mods:date>
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<mods:number id="96FDE30D8545D3872EE659E338A30B21">2004-01-01</mods:number>
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<mods:number id="719673CE87E5645A7CC943EA4BCC4EED">38</mods:number>
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<mods:number id="44FD49CCB54C36A626C801C39FEF6A1C">11</mods:number>
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<mods:classification id="0DD84E12F29B7E560E1DD1CE0881AFA1">journal article</mods:classification>
<mods:identifier id="B01A28C45B5E484C78CEE5D1C8964064" type="DOI">10.1080/0022293031000155395</mods:identifier>
<mods:identifier id="38051C7CC1DB9D8A7FCEF119D8514387" type="ISSN">1464-5262</mods:identifier>
<mods:identifier id="09FBB4A8061D47F3A3FE4BCADBA3AE46" type="Zenodo-Dep">5250248</mods:identifier>
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<treatment id="537A7F7AFF94FD2B564E9B71707DFBE9" LSID="urn:lsid:plazi:treatment:537A7F7AFF94FD2B564E9B71707DFBE9" httpUri="http://treatment.plazi.org/id/537A7F7AFF94FD2B564E9B71707DFBE9" lastPageId="10" lastPageNumber="1348" pageId="4" pageNumber="1342">
<subSubSection id="93C99DE7FF94FD25564E9B71774AFBFE" box="[513,864,1047,1071]" pageId="4" pageNumber="1342" type="nomenclature">
<paragraph id="DB6CCE6CFF94FD25564E9B71774AFBFE" blockId="4.[513,864,1047,1071]" box="[513,864,1047,1071]" pageId="4" pageNumber="1342">
<heading id="80247900FF94FD25564E9B71774AFBFE" box="[513,864,1047,1071]" centered="true" fontSize="10" level="2" pageId="4" pageNumber="1342" reason="5">
<taxonomicName id="1CD3B5EFFF94FD25564E9B717724FBFE" ID-CoL="77SG6" authority="Williams, 2004" authorityName="Williams" authorityYear="2004" box="[513,782,1047,1071]" class="Polychaeta" family="Spionidae" genus="Polydorella" kingdom="Animalia" order="Spionida" pageId="4" pageNumber="1342" phylum="Annelida" rank="species" species="kamakamai" status="sp. nov.">
<emphasis id="E9A7127EFF94FD25564E9B717724FBFE" bold="true" box="[513,782,1047,1071]" italics="true" pageId="4" pageNumber="1342">Polydorella kamakamai</emphasis>
</taxonomicName>
,
<taxonomicNameLabel id="F294AF05FF94FD25576F9B71774AFBFE" box="[800,864,1047,1071]" pageId="4" pageNumber="1342" rank="species">sp. n.</taxonomicNameLabel>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="93C99DE7FF94FD2556119B5176CDFB9E" box="[606,743,1079,1103]" pageId="4" pageNumber="1342" type="description">
<paragraph id="DB6CCE6CFF94FD2556119B5176CDFB9E" blockId="4.[606,743,1079,1103]" box="[606,743,1079,1103]" pageId="4" pageNumber="1342">
(
<figureCitation id="43E8D2E9FF94FD25562A9B5176F4FB9E" box="[613,734,1079,1103]" captionStart-0="FIG" captionStart-1="FIG" captionStart-2="FIG" captionStart-3="FIG" captionStart-4="FIG" captionStartId-0="3.[159,175,1286,1308]" captionStartId-1="5.[159,175,1364,1386]" captionStartId-2="6.[154,170,1594,1616]" captionStartId-3="7.[159,175,875,897]" captionStartId-4="8.[154,170,871,893]" captionTargetBox-0="[159,1115,172,1246]" captionTargetBox-1="[159,1116,171,1321]" captionTargetBox-2="[161,1109,211,1550]" captionTargetBox-3="[309,969,173,834]" captionTargetBox-4="[157,1109,175,830]" captionTargetId-0="figure-98@3.[159,1117,170,1248]" captionTargetId-1="figure-51@5.[159,1116,170,1325]" captionTargetId-2="figure-1@6.[154,1112,207,1555]" captionTargetId-3="figure-255@7.[306,970,170,837]" captionTargetId-4="figure-173@8.[154,1112,170,833]" captionTargetPageId-0="3" captionTargetPageId-1="5" captionTargetPageId-2="6" captionTargetPageId-3="7" captionTargetPageId-4="8" captionText-0="FIG. 1. Polydorella kamakamai new species: holotype (USNM 1008710) and paratypes (AMNH 4347). (A) Dorsal view of paratype exhibiting sexual reproduction (developing eggs in segments 13–15 outlined in dotted lines; three single eggs shown via shading); (B) dorsal view of holotype, arrowhead indicating growth zone following segment 10; (C) capillary neurosetae from segment 4; (D) spines of fifth segment, ventral view; (E) four acicular and two capillary neurosetae from segment 4; (F) hooded hook from segment 8. Scale: 100 mm (A, B); 10mm (C–F)." captionText-1="FIG. 2. Polydorella kamakamai new species: paratypes (USNM 1008712), SEM micrographs. (A) Spines of fifth segment, lateral view; (B) magnified view of spines in (A), showing digitiform bosses on ventral row of spines; (C) dorsal view of individual with developing stolon 1; (D) dorsal view of paratomic chain with stock, stolon 2 (St2) and stolon 1 (St1); (E) lateral view of paratomic chain with stock, stolon 2, stolon 1 and developing secondary stolon (2‡St). Scale: 5 mm (A); 1 mm (B); 100 mm (C–E)." captionText-2="FIG. 3. Polydorella kamakamai new species: paratype (USNM 1008711). Paratomic chain with five developing individuals. (A) Dorsal view; (B) ventral view. Segment numbers of stock, stolon 2 and stolon 1 indicated to left of ventral view. P, pygidium. Scale: 250 mm." captionText-3="FIG. 4. Polydorella kamakamai new species. Percentages of single individuals and paratomic chains of two, three, four and five or more individuals in colonies from the Philippines (N~290)." captionText-4="FIG. 5. Polydorella kamakamai new species. Semi-diagrammatic view of process of asexual reproduction via paratomy. (A) Single stock individual without growth zone (left palp shown for comparison with developing palp buds; palps removed from remaining figures); (B) development of stolon 1 (St1) growth zone; (C) development of stolon 2 (St2) growth zone; (D) development of first 10 segments of stolon 1 and stolon 3 (St3) growth zone; secondary stolon (2‡St) indicated by arrowhead; (E) paratomic chain of five developing individuals just prior to paratomic division between stolon 2 and stolon 1; site of paratomic division indicated by arrow. Scale: 250 mm." figureDoi-0="http://doi.org/10.5281/zenodo.5250250" figureDoi-1="http://doi.org/10.5281/zenodo.5250252" figureDoi-2="http://doi.org/10.5281/zenodo.5250254" figureDoi-3="http://doi.org/10.5281/zenodo.5250256" figureDoi-4="http://doi.org/10.5281/zenodo.5250258" httpUri-0="https://zenodo.org/record/5250250/files/figure.png" httpUri-1="https://zenodo.org/record/5250252/files/figure.png" httpUri-2="https://zenodo.org/record/5250254/files/figure.png" httpUri-3="https://zenodo.org/record/5250256/files/figure.png" httpUri-4="https://zenodo.org/record/5250258/files/figure.png" pageId="4" pageNumber="1342">figures 1–5</figureCitation>
)
</paragraph>
</subSubSection>
<subSubSection id="93C99DE7FF94FD2B548D9B14707DFBE9" lastPageId="10" lastPageNumber="1348" pageId="4" pageNumber="1342" type="materials_examined">
<paragraph id="DB6CCE6CFF94FD25548D9B1477BBFB7B" blockId="4.[154,1111,1138,1450]" pageId="4" pageNumber="1342">
<materialsCitation id="6BBBC431FF94FD25548D9B1477BBFB7B" collectingDate="2000-07-01" collectionCode="USNM" country="Philippines" elevation="3" location="Puerto Galera" municipality="Agus Beach" pageId="4" pageNumber="1342" specimenCode="USNM 1008710" specimenCount="1" typeStatus="holotype">
<typeStatus id="046870CEFF94FD25548D9B147510FB59" box="[194,314,1138,1162]" pageId="4" pageNumber="1342" type="holotype">HOLOTYPE</typeStatus>
:
<collectingCountry id="A3C48EFCFF94FD2555029B1475E4FB5B" box="[333,462,1138,1162]" name="Philippines" pageId="4" pageNumber="1342">Philippines</collectingCountry>
,
<location id="DE0C98B7FF94FD2555919B1476ADFB5B" LSID="urn:lsid:plazi:treatment:537A7F7AFF94FD2B564E9B71707DFBE9:DE0C98B7FF94FD2555919B1476ADFB5B" box="[478,647,1138,1162]" country="Philippines" municipality="Agus Beach" name="Puerto Galera" pageId="4" pageNumber="1342">Puerto Galera</location>
,
<collectingMunicipality id="3B085416FF94FD2556D79B14770DFB5B" box="[664,807,1138,1162]" pageId="4" pageNumber="1342">Agus Beach</collectingMunicipality>
,
<emphasis id="E9A7127EFF94FD2557649B14776CFB58" bold="true" box="[811,838,1138,1161]" italics="true" pageId="4" pageNumber="1342">y</emphasis>
<quantity id="1C2B6389FF94FD2557099B14775BFB5B" box="[838,881,1138,1162]" metricMagnitude="0" metricUnit="m" metricValue="3.0" pageId="4" pageNumber="1342" unit="m" value="3.0">
<elevation id="50FE295FFF94FD2557099B14775BFB5B" box="[838,881,1138,1162]" metricMagnitude="0" metricUnit="m" metricValue="3.0" pageId="4" pageNumber="1342" unit="m" value="3.0">3 m</elevation>
</quantity>
,
<date id="AF6DE8ACFF94FD2557CE9B147024FB5B" box="[897,1038,1138,1162]" pageId="4" pageNumber="1342" value="2000-07-01">
<collectingDate id="BF291144FF94FD2557CE9B147024FB5B" box="[897,1038,1138,1162]" pageId="4" pageNumber="1342" value="2000-07-01">1 July 2000</collectingDate>
</date>
, from
<taxonomicName id="1CD3B5EFFF94FD2554D59BF4769BFB7B" authority="(Thiele, 1903)" baseAuthorityName="Thiele" baseAuthorityYear="1903" box="[154,689,1170,1194]" class="Demospongiae" family="Microcionidae" genus="Clathria" kingdom="Animalia" order="Poecilosclerida" pageId="4" pageNumber="1342" phylum="Porifera" rank="species" species="cervicornis" subGenus="Thalysias">
<emphasis id="E9A7127EFF94FD2554D59BF4762CFB7B" box="[154,518,1170,1194]" italics="true" pageId="4" pageNumber="1342">Clathria (Thalysias) cervicornis</emphasis>
(Thiele, 1903)
</taxonomicName>
(
<specimenCode id="8B756617FF94FD25568C9BF477AFFB7B" box="[707,901,1170,1194]" collectionCode="USNM" country="USA" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34871" lsid="urn:lsid:biocol.org:col:34871" name="Smithsonian Institution, National Museum of Natural History" pageId="4" pageNumber="1342" type="Museum">USNM 1008710</specimenCode>
).
</materialsCitation>
</paragraph>
<paragraph id="DB6CCE6CFF94FD25548D9BD47631FAFB" blockId="4.[154,1111,1138,1450]" pageId="4" pageNumber="1342">
<materialsCitation id="6BBBC431FF94FD25548D9BD4763DFAFB" collectingDate="2000-07-24" collectionCode="USNM, AMNH" country="Philippines" county="Puerto Galera" elevation="3" location="Beach" municipality="Small" pageId="4" pageNumber="1342" specimenCode="USNM 1008711, USNM 1008712, AMNH 4347" specimenCount="10" stateProvince="Laguna" typeStatus="paratype">
<typeStatus id="046870CEFF94FD25548D9BD4756BFB19" box="[194,321,1202,1226]" pageId="4" pageNumber="1342" type="paratype">PARATYPES</typeStatus>
: same data as for holotype (
<specimenCode id="8B756617FF94FD2556F99BD47756FB1B" box="[694,892,1202,1226]" collectionCode="USNM" country="USA" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34871" lsid="urn:lsid:biocol.org:col:34871" name="Smithsonian Institution, National Museum of Natural History" pageId="4" pageNumber="1342" type="Museum">USNM 1008711</specimenCode>
: 270z
<typeStatus id="046870CEFF94FD2557AE9BD47078FB1B" box="[993,1106,1202,1226]" pageId="4" pageNumber="1342" type="paratype">paratypes</typeStatus>
;
<specimenCode id="8B756617FF94FD2554D59BB47576FB3B" box="[154,348,1234,1258]" collectionCode="USNM" country="USA" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34871" lsid="urn:lsid:biocol.org:col:34871" name="Smithsonian Institution, National Museum of Natural History" pageId="4" pageNumber="1342" type="Museum">USNM 1008712</specimenCode>
:
<specimenCount id="CDD505E5FF94FD2555239BB47629FB3B" box="[364,515,1234,1258]" pageId="4" pageNumber="1342" type="generic" typeStatus="paratypes">10 paratypes</specimenCount>
, on nine SEM stubs).
<collectingCountry id="A3C48EFCFF94FD2557579BB477B3FB3B" box="[792,921,1234,1258]" name="Philippines" pageId="4" pageNumber="1342">Philippines</collectingCountry>
,
<collectingCounty id="320DB6E0FF94FD2557E69BB47078FB3B" box="[937,1106,1234,1258]" pageId="4" pageNumber="1342">Puerto Galera</collectingCounty>
,
<collectingMunicipality id="3B085416FF94FD2554D59B9474F0FADB" box="[154,218,1266,1290]" pageId="4" pageNumber="1342">Small</collectingMunicipality>
<collectingRegion id="1917008EFF94FD2554AC9B947549FADB" box="[227,355,1266,1290]" country="Philippines" name="Laguna" pageId="4" pageNumber="1342">La Laguna</collectingRegion>
<location id="DE0C98B7FF94FD2555239B94759EFADB" LSID="urn:lsid:plazi:treatment:537A7F7AFF94FD2B564E9B71707DFBE9:DE0C98B7FF94FD2555239B94759EFADB" box="[364,436,1266,1290]" country="Philippines" county="Puerto Galera" municipality="Small" name="Beach" pageId="4" pageNumber="1342" stateProvince="Laguna">Beach</location>
,
<emphasis id="E9A7127EFF94FD2555F79B9475F9FAD8" bold="true" box="[440,467,1266,1289]" italics="true" pageId="4" pageNumber="1342">y</emphasis>
<quantity id="1C2B6389FF94FD25559C9B9475D7FADB" box="[467,509,1266,1290]" metricMagnitude="0" metricUnit="m" metricValue="3.0" pageId="4" pageNumber="1342" unit="m" value="3.0">
<elevation id="50FE295FFF94FD25559C9B9475D7FADB" box="[467,509,1266,1290]" metricMagnitude="0" metricUnit="m" metricValue="3.0" pageId="4" pageNumber="1342" unit="m" value="3.0">3 m</elevation>
</quantity>
,
<date id="AF6DE8ACFF94FD2556449B9476B4FADB" box="[523,670,1266,1290]" pageId="4" pageNumber="1342" value="2000-07-24">
<collectingDate id="BF291144FF94FD2556449B9476B4FADB" box="[523,670,1266,1290]" pageId="4" pageNumber="1342" value="2000-07-24">24 July 2000</collectingDate>
</date>
, from
<taxonomicName id="1CD3B5EFFF94FD2556A29B94707CFADB" baseAuthorityName="Thiele" baseAuthorityYear="1903" box="[749,1110,1266,1290]" class="Demospongiae" family="Microcionidae" genus="Clathria" kingdom="Animalia" order="Poecilosclerida" pageId="4" pageNumber="1342" phylum="Porifera" rank="species" species="cervicornis" subGenus="Thalysias">
<emphasis id="E9A7127EFF94FD2556A29B94707CFADB" box="[749,1110,1266,1290]" italics="true" pageId="4" pageNumber="1342">Clathria (Thalysias) cervicornis</emphasis>
</taxonomicName>
(
<specimenCode id="8B756617FF94FD2554ED9A74756BFAFB" box="[162,321,1298,1322]" collectionCode="AMNH" country="USA" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34925" lsid="urn:lsid:biocol.org:col:34925" name="American Museum of Natural History" pageId="4" pageNumber="1342">AMNH 4347</specimenCode>
: 180z
<typeStatus id="046870CEFF94FD2555D39A747625FAFB" box="[412,527,1298,1322]" pageId="4" pageNumber="1342" type="paratype">paratypes</typeStatus>
)
</materialsCitation>
.
</paragraph>
<paragraph id="DB6CCE6CFF94FD25548D9A547779FA7B" blockId="4.[154,1111,1138,1450]" pageId="4" pageNumber="1342">
<emphasis id="E9A7127EFF94FD25548D9A547568FA9B" box="[194,322,1330,1354]" italics="true" pageId="4" pageNumber="1342">Etymology.</emphasis>
The specific name
<taxonomicName id="1CD3B5EFFF94FD25567E9A547699FA9B" authorityName="Williams" authorityYear="2004" box="[561,691,1330,1354]" class="Polychaeta" family="Spionidae" genus="Polydorella" kingdom="Animalia" order="Spionida" pageId="4" pageNumber="1342" phylum="Annelida" rank="species" species="kamakamai">
<emphasis id="E9A7127EFF94FD25567E9A547699FA9B" box="[561,691,1330,1354]" italics="true" pageId="4" pageNumber="1342">kamakamai</emphasis>
</taxonomicName>
is derived from the Filipino word for supernatural beings that are small, human-like creatures possessing long beards and nails used to pinch offending children (
<bibRefCitation id="BF42B39DFF94FD2556B39A1477B3FA5B" author="JOCANO, F. L." box="[764,921,1394,1418]" pageId="4" pageNumber="1342" refId="ref8892" refString="JOCANO, F. L., 1969, Growing up in a Philippine barrio (New York: Holt, Rinehart and Winston), 121 pp." type="book" year="1969">Jocano, 1969</bibRefCitation>
), reminiscent of the acicular neurosetae of anterior segments in this species.
</paragraph>
<paragraph id="DB6CCE6CFF94FD2554D59A8B7533F9D4" blockId="4.[154,1111,1517,1732]" box="[154,281,1517,1541]" pageId="4" pageNumber="1342">
<emphasis id="E9A7127EFF94FD2554D59A8B7533F9D4" box="[154,281,1517,1541]" italics="true" pageId="4" pageNumber="1342">Description</emphasis>
</paragraph>
<paragraph id="DB6CCE6CFF94FD25548D996A76FEF915" blockId="4.[154,1111,1517,1732]" pageId="4" pageNumber="1342">
<emphasis id="E9A7127EFF94FD25548D996A751AF9F5" box="[194,304,1548,1572]" italics="true" pageId="4" pageNumber="1342">
<typeStatus id="046870CEFF94FD25548D996A7507F9F5" box="[194,301,1548,1572]" pageId="4" pageNumber="1342" type="holotype">Holotype</typeStatus>
.
</emphasis>
1.3 mm long, 0.3 mm wide at segment 7; 14 segments. Prostomium bifid; caruncle short, terminating between posterior margin of segment 1 and middle of segment 2; occipital tentacle and eyes absent (
<figureCitation id="43E8D2E9FF94FD255732992A77D2F9B5" box="[893,1016,1612,1636]" captionStart-0="FIG" captionStart-1="FIG" captionStart-2="FIG" captionStartId-0="3.[159,175,1286,1308]" captionStartId-1="5.[159,175,1364,1386]" captionStartId-2="6.[154,170,1594,1616]" captionTargetBox-0="[159,1115,172,1246]" captionTargetBox-1="[159,1116,171,1321]" captionTargetBox-2="[161,1109,211,1550]" captionTargetId-0="figure-98@3.[159,1117,170,1248]" captionTargetId-1="figure-51@5.[159,1116,170,1325]" captionTargetId-2="figure-1@6.[154,1112,207,1555]" captionTargetPageId-0="3" captionTargetPageId-1="5" captionTargetPageId-2="6" captionText-0="FIG. 1. Polydorella kamakamai new species: holotype (USNM 1008710) and paratypes (AMNH 4347). (A) Dorsal view of paratype exhibiting sexual reproduction (developing eggs in segments 13–15 outlined in dotted lines; three single eggs shown via shading); (B) dorsal view of holotype, arrowhead indicating growth zone following segment 10; (C) capillary neurosetae from segment 4; (D) spines of fifth segment, ventral view; (E) four acicular and two capillary neurosetae from segment 4; (F) hooded hook from segment 8. Scale: 100 mm (A, B); 10mm (C–F)." captionText-1="FIG. 2. Polydorella kamakamai new species: paratypes (USNM 1008712), SEM micrographs. (A) Spines of fifth segment, lateral view; (B) magnified view of spines in (A), showing digitiform bosses on ventral row of spines; (C) dorsal view of individual with developing stolon 1; (D) dorsal view of paratomic chain with stock, stolon 2 (St2) and stolon 1 (St1); (E) lateral view of paratomic chain with stock, stolon 2, stolon 1 and developing secondary stolon (2‡St). Scale: 5 mm (A); 1 mm (B); 100 mm (C–E)." captionText-2="FIG. 3. Polydorella kamakamai new species: paratype (USNM 1008711). Paratomic chain with five developing individuals. (A) Dorsal view; (B) ventral view. Segment numbers of stock, stolon 2 and stolon 1 indicated to left of ventral view. P, pygidium. Scale: 250 mm." figureDoi-0="http://doi.org/10.5281/zenodo.5250250" figureDoi-1="http://doi.org/10.5281/zenodo.5250252" figureDoi-2="http://doi.org/10.5281/zenodo.5250254" httpUri-0="https://zenodo.org/record/5250250/files/figure.png" httpUri-1="https://zenodo.org/record/5250252/files/figure.png" httpUri-2="https://zenodo.org/record/5250254/files/figure.png" pageId="4" pageNumber="1342">figures 1–3</figureCitation>
). Palps extend posteriorly for 10–12 segments. Black pigmentation in irregular patches on palps, sides of peristomium, prostomium, along caruncle, dorsal and ventral sides of all segments and pygidium (
<figureCitation id="43E8D2E9FF94FD25560899CA768CF915" box="[583,678,1708,1732]" captionStart="FIG" captionStartId="3.[159,175,1286,1308]" captionTargetBox="[159,1115,172,1246]" captionTargetId="figure-98@3.[159,1117,170,1248]" captionTargetPageId="3" captionText="FIG. 1. Polydorella kamakamai new species: holotype (USNM 1008710) and paratypes (AMNH 4347). (A) Dorsal view of paratype exhibiting sexual reproduction (developing eggs in segments 13–15 outlined in dotted lines; three single eggs shown via shading); (B) dorsal view of holotype, arrowhead indicating growth zone following segment 10; (C) capillary neurosetae from segment 4; (D) spines of fifth segment, ventral view; (E) four acicular and two capillary neurosetae from segment 4; (F) hooded hook from segment 8. Scale: 100 mm (A, B); 10mm (C–F)." figureDoi="http://doi.org/10.5281/zenodo.5250250" httpUri="https://zenodo.org/record/5250250/files/figure.png" pageId="4" pageNumber="1342">figures 1</figureCitation>
,
<figureCitation id="43E8D2E9FF94FD2556F899CA76ECF915" box="[695,710,1708,1732]" captionStart="FIG" captionStartId="6.[154,170,1594,1616]" captionTargetBox="[161,1109,211,1550]" captionTargetId="figure-1@6.[154,1112,207,1555]" captionTargetPageId="6" captionText="FIG. 3. Polydorella kamakamai new species: paratype (USNM 1008711). Paratomic chain with five developing individuals. (A) Dorsal view; (B) ventral view. Segment numbers of stock, stolon 2 and stolon 1 indicated to left of ventral view. P, pygidium. Scale: 250 mm." figureDoi="http://doi.org/10.5281/zenodo.5250254" httpUri="https://zenodo.org/record/5250254/files/figure.png" pageId="4" pageNumber="1342">3</figureCitation>
).
</paragraph>
<caption id="8FAC9EE4FF95FD2454D09A327039FA3F" ID-DOI="http://doi.org/10.5281/zenodo.5250252" ID-Zenodo-Dep="5250252" httpUri="https://zenodo.org/record/5250252/files/figure.png" pageId="5" pageNumber="1343" startId="5.[159,175,1364,1386]" targetBox="[159,1116,171,1321]" targetPageId="5">
<paragraph id="DB6CCE6CFF95FD2454D09A327039FA3F" blockId="5.[159,1116,1364,1518]" pageId="5" pageNumber="1343">
FIG. 2.
<taxonomicName id="1CD3B5EFFF95FD2454B69A3275C4FAB8" authority="Williams, 2004" authorityName="Williams" authorityYear="2004" box="[249,494,1364,1385]" class="Polychaeta" family="Spionidae" genus="Polydorella" kingdom="Animalia" order="Spionida" pageId="5" pageNumber="1343" phylum="Annelida" rank="species" species="kamakamai" status="sp. nov.">
<emphasis id="E9A7127EFF95FD2454B69A3275C4FAB8" box="[249,494,1364,1385]" italics="true" pageId="5" pageNumber="1343">Polydorella kamakamai</emphasis>
</taxonomicName>
<taxonomicNameLabel id="F294AF05FF95FD2455B19A327654FABB" box="[510,638,1364,1386]" pageId="5" pageNumber="1343" rank="species">new species</taxonomicNameLabel>
: paratypes (USNM 1008712), SEM micrographs. (A) Spines of fifth segment, lateral view; (B) magnified view of spines in (A), showing digitiform bosses on ventral row of spines; (C) dorsal view of individual with developing stolon 1; (D) dorsal view of paratomic chain with stock, stolon 2 (St2) and stolon 1 (St1); (E) lateral view of paratomic chain with stock, stolon 2, stolon 1 and developing secondary stolon (2
<emphasis id="E9A7127EFF95FD2456659ABE7619FA3F" bold="true" box="[554,563,1496,1518]" pageId="5" pageNumber="1343">‡</emphasis>
St). Scale: 5
<emphasis id="E9A7127EFF95FD2456FA9ABF76E8FA3F" bold="true" box="[693,706,1497,1518]" italics="true" pageId="5" pageNumber="1343">m</emphasis>
m (A); 1
<emphasis id="E9A7127EFF95FD24576D9ABF7705FA3F" bold="true" box="[802,815,1497,1518]" italics="true" pageId="5" pageNumber="1343">m</emphasis>
m (B); 100
<emphasis id="E9A7127EFF95FD2457EB9ABF779BFA3F" bold="true" box="[932,945,1497,1518]" italics="true" pageId="5" pageNumber="1343">m</emphasis>
m (C–E).
</paragraph>
</caption>
<paragraph id="DB6CCE6CFF95FD26548899257536FB4B" blockId="5.[159,1116,1603,1729]" lastBlockId="7.[159,1116,989,1732]" lastPageId="7" lastPageNumber="1345" pageId="5" pageNumber="1343">
Segment 1 without notopodial lobes or notosetae; neuropodia lobes present and neurosetae absent. Capillary notosetae of segments 2–4, 6 and subsequent segments in two rows; no specialized posterior notosetae. Neurosetae of segments 2–5 of
<specimenCount id="CDD505E5FF95FD24507E99E174F7F910" pageId="5" pageNumber="1343" type="generic" typeStatus="types">two types</specimenCount>
, arranged in two rows; dorsal row composed of up to three acicular spines on segment four, ventral row composed of up to four capillaries on segment four (
<figureCitation id="43E8D2E9FF97FD2654E89C987512FBC7" box="[167,312,1022,1046]" captionStart="FIG" captionStartId="3.[159,175,1286,1308]" captionTargetBox="[159,1115,172,1246]" captionTargetId="figure-98@3.[159,1117,170,1248]" captionTargetPageId="3" captionText="FIG. 1. Polydorella kamakamai new species: holotype (USNM 1008710) and paratypes (AMNH 4347). (A) Dorsal view of paratype exhibiting sexual reproduction (developing eggs in segments 13–15 outlined in dotted lines; three single eggs shown via shading); (B) dorsal view of holotype, arrowhead indicating growth zone following segment 10; (C) capillary neurosetae from segment 4; (D) spines of fifth segment, ventral view; (E) four acicular and two capillary neurosetae from segment 4; (F) hooded hook from segment 8. Scale: 100 mm (A, B); 10mm (C–F)." figureDoi="http://doi.org/10.5281/zenodo.5250250" httpUri="https://zenodo.org/record/5250250/files/figure.png" pageId="7" pageNumber="1345">figure 1C, E</figureCitation>
); segments 6 and 7 with reduced number of acicular spines and capillaries; three bidentate hooded hooks begin on segment 8, not accompanied by capillaries; hooks with approximately right angle between main fang and shaft, narrow angle between main fang and apical tooth, with constriction on shaft (
<figureCitation id="43E8D2E9FF97FD2654E89BE47524FB4B" box="[167,270,1154,1178]" captionStart="FIG" captionStartId="3.[159,175,1286,1308]" captionTargetBox="[159,1115,172,1246]" captionTargetId="figure-98@3.[159,1117,170,1248]" captionTargetPageId="3" captionText="FIG. 1. Polydorella kamakamai new species: holotype (USNM 1008710) and paratypes (AMNH 4347). (A) Dorsal view of paratype exhibiting sexual reproduction (developing eggs in segments 13–15 outlined in dotted lines; three single eggs shown via shading); (B) dorsal view of holotype, arrowhead indicating growth zone following segment 10; (C) capillary neurosetae from segment 4; (D) spines of fifth segment, ventral view; (E) four acicular and two capillary neurosetae from segment 4; (F) hooded hook from segment 8. Scale: 100 mm (A, B); 10mm (C–F)." figureDoi="http://doi.org/10.5281/zenodo.5250250" httpUri="https://zenodo.org/record/5250250/files/figure.png" pageId="7" pageNumber="1345">figure 1F</figureCitation>
).
</paragraph>
<caption id="8FAC9EE4FF96FD2754D5995C7505F94E" ID-DOI="http://doi.org/10.5281/zenodo.5250254" ID-Zenodo-Dep="5250254" httpUri="https://zenodo.org/record/5250254/files/figure.png" pageId="6" pageNumber="1344" startId="6.[154,170,1594,1616]" targetBox="[161,1109,211,1550]" targetPageId="6">
<paragraph id="DB6CCE6CFF96FD2754D5995C7505F94E" blockId="6.[154,1111,1594,1695]" pageId="6" pageNumber="1344">
FIG. 3.
<taxonomicName id="1CD3B5EFFF96FD2754BB995C75CFF99E" authority="Williams, 2004" authorityName="Williams" authorityYear="2004" box="[244,485,1594,1615]" class="Polychaeta" family="Spionidae" genus="Polydorella" kingdom="Animalia" order="Spionida" pageId="6" pageNumber="1344" phylum="Annelida" rank="species" species="kamakamai" status="sp. nov.">
<emphasis id="E9A7127EFF96FD2754BB995C75CFF99E" box="[244,485,1594,1615]" italics="true" pageId="6" pageNumber="1344">Polydorella kamakamai</emphasis>
</taxonomicName>
<taxonomicNameLabel id="F294AF05FF96FD2755BF995C7647F981" box="[496,621,1594,1616]" pageId="6" pageNumber="1344" rank="species">new species</taxonomicNameLabel>
: paratype (USNM 1008711). Paratomic chain with five developing individuals. (A) Dorsal view; (B) ventral view. Segment numbers of stock, stolon 2 and stolon 1 indicated to left of ventral view. P, pygidium. Scale: 250
<emphasis id="E9A7127EFF96FD27554899EF753EF94F" bold="true" box="[263,276,1673,1694]" italics="true" pageId="6" pageNumber="1344">m</emphasis>
m.
</paragraph>
</caption>
<caption id="8FAC9EE4FF97FD2654D09C0D7560FC67" ID-DOI="http://doi.org/10.5281/zenodo.5250256" ID-Zenodo-Dep="5250256" httpUri="https://zenodo.org/record/5250256/files/figure.png" pageId="7" pageNumber="1345" startId="7.[159,175,875,897]" targetBox="[309,969,173,834]" targetPageId="7">
<paragraph id="DB6CCE6CFF97FD2654D09C0D7560FC67" blockId="7.[159,1116,875,950]" pageId="7" pageNumber="1345">
FIG. 4.
<taxonomicName id="1CD3B5EFFF97FD2654B69C0A75CDFC50" authority="Williams, 2004" authorityName="Williams" authorityYear="2004" box="[249,487,876,897]" class="Polychaeta" family="Spionidae" genus="Polydorella" kingdom="Animalia" order="Spionida" pageId="7" pageNumber="1345" phylum="Annelida" rank="species" species="kamakamai" status="sp. nov.">
<emphasis id="E9A7127EFF97FD2654B69C0A75CDFC50" box="[249,487,876,897]" italics="true" pageId="7" pageNumber="1345">Polydorella kamakamai</emphasis>
</taxonomicName>
<taxonomicNameLabel id="F294AF05FF97FD2655A09C0D7642FC50" box="[495,616,875,897]" pageId="7" pageNumber="1345" rank="species">new species</taxonomicNameLabel>
. Percentages of single individuals and paratomic chains of two, three, four and five or more individuals in colonies from the Philippines (
<emphasis id="E9A7127EFF97FD2654A39CC774D4FC67" box="[236,254,929,950]" italics="true" pageId="7" pageNumber="1345">N</emphasis>
~290).
</paragraph>
</caption>
<paragraph id="DB6CCE6CFF97FD2654889BC475FBFA51" blockId="7.[159,1116,989,1732]" pageId="7" pageNumber="1345">
Segment 5 slightly modified, about the same size as segments 4 and 6, with acicular spines and capillary neurosetae in same arrangement as on segments 2–4; notosetae absent (
<figureCitation id="43E8D2E9FF97FD2655CA9B8275DBFB2D" box="[389,497,1252,1276]" captionStart="FIG" captionStartId="5.[159,175,1364,1386]" captionTargetBox="[159,1116,171,1321]" captionTargetId="figure-51@5.[159,1116,170,1325]" captionTargetPageId="5" captionText="FIG. 2. Polydorella kamakamai new species: paratypes (USNM 1008712), SEM micrographs. (A) Spines of fifth segment, lateral view; (B) magnified view of spines in (A), showing digitiform bosses on ventral row of spines; (C) dorsal view of individual with developing stolon 1; (D) dorsal view of paratomic chain with stock, stolon 2 (St2) and stolon 1 (St1); (E) lateral view of paratomic chain with stock, stolon 2, stolon 1 and developing secondary stolon (2‡St). Scale: 5 mm (A); 1 mm (B); 100 mm (C–E)." figureDoi="http://doi.org/10.5281/zenodo.5250252" httpUri="https://zenodo.org/record/5250252/files/figure.png" pageId="7" pageNumber="1345">figure 2A</figureCitation>
). Two
<typeStatus id="046870CEFF97FD2656169B8276BEFB2D" box="[601,660,1252,1276]" pageId="7" pageNumber="1345">types</typeStatus>
of major spines on segment
<quantity id="1C2B6389FF97FD26506A9B827076FB2D" box="[1061,1116,1252,1276]" metricMagnitude="-1" metricUnit="m" metricValue="1.27" pageId="7" pageNumber="1345" unit="in" value="5.0">5 in</quantity>
approximately straight, horizontal rows; ventral row of four acicular spines with small digitiform bosses (
<emphasis id="E9A7127EFF97FD2655F39A4075FDFAEC" bold="true" box="[444,471,1318,1341]" italics="true" pageId="7" pageNumber="1345">y</emphasis>
0.3
<emphasis id="E9A7127EFF97FD2655B29A407626FAEF" bold="true" box="[509,524,1318,1342]" italics="true" pageId="7" pageNumber="1345">m</emphasis>
m) closely applied to the apical ends of the spines, lacking on tips of the spines (
<figureCitation id="43E8D2E9FF97FD2655B39A217658FA8E" box="[508,626,1351,1375]" captionStart="FIG" captionStartId="3.[159,175,1286,1308]" captionTargetBox="[159,1115,172,1246]" captionTargetId="figure-98@3.[159,1117,170,1248]" captionTargetPageId="3" captionText="FIG. 1. Polydorella kamakamai new species: holotype (USNM 1008710) and paratypes (AMNH 4347). (A) Dorsal view of paratype exhibiting sexual reproduction (developing eggs in segments 13–15 outlined in dotted lines; three single eggs shown via shading); (B) dorsal view of holotype, arrowhead indicating growth zone following segment 10; (C) capillary neurosetae from segment 4; (D) spines of fifth segment, ventral view; (E) four acicular and two capillary neurosetae from segment 4; (F) hooded hook from segment 8. Scale: 100 mm (A, B); 10mm (C–F)." figureDoi="http://doi.org/10.5281/zenodo.5250250" httpUri="https://zenodo.org/record/5250250/files/figure.png" pageId="7" pageNumber="1345">figures 1D</figureCitation>
,
<figureCitation id="43E8D2E9FF97FD2656CD9A2176E3FA8E" box="[642,713,1351,1375]" captionStart="FIG" captionStartId="5.[159,175,1364,1386]" captionTargetBox="[159,1116,171,1321]" captionTargetId="figure-51@5.[159,1116,170,1325]" captionTargetPageId="5" captionText="FIG. 2. Polydorella kamakamai new species: paratypes (USNM 1008712), SEM micrographs. (A) Spines of fifth segment, lateral view; (B) magnified view of spines in (A), showing digitiform bosses on ventral row of spines; (C) dorsal view of individual with developing stolon 1; (D) dorsal view of paratomic chain with stock, stolon 2 (St2) and stolon 1 (St1); (E) lateral view of paratomic chain with stock, stolon 2, stolon 1 and developing secondary stolon (2‡St). Scale: 5 mm (A); 1 mm (B); 100 mm (C–E)." figureDoi="http://doi.org/10.5281/zenodo.5250252" httpUri="https://zenodo.org/record/5250252/files/figure.png" pageId="7" pageNumber="1345">2A, B</figureCitation>
), and a dorsal row of four falcate spines (
<figureCitation id="43E8D2E9FF97FD2654B99A0E7546FA51" box="[246,364,1384,1408]" captionStart="FIG" captionStartId="3.[159,175,1286,1308]" captionTargetBox="[159,1115,172,1246]" captionTargetId="figure-98@3.[159,1117,170,1248]" captionTargetPageId="3" captionText="FIG. 1. Polydorella kamakamai new species: holotype (USNM 1008710) and paratypes (AMNH 4347). (A) Dorsal view of paratype exhibiting sexual reproduction (developing eggs in segments 13–15 outlined in dotted lines; three single eggs shown via shading); (B) dorsal view of holotype, arrowhead indicating growth zone following segment 10; (C) capillary neurosetae from segment 4; (D) spines of fifth segment, ventral view; (E) four acicular and two capillary neurosetae from segment 4; (F) hooded hook from segment 8. Scale: 100 mm (A, B); 10mm (C–F)." figureDoi="http://doi.org/10.5281/zenodo.5250250" httpUri="https://zenodo.org/record/5250250/files/figure.png" pageId="7" pageNumber="1345">figures 1D</figureCitation>
,
<figureCitation id="43E8D2E9FF97FD2655339A0E75EFFA51" box="[380,453,1384,1408]" captionStart="FIG" captionStartId="5.[159,175,1364,1386]" captionTargetBox="[159,1116,171,1321]" captionTargetId="figure-51@5.[159,1116,170,1325]" captionTargetPageId="5" captionText="FIG. 2. Polydorella kamakamai new species: paratypes (USNM 1008712), SEM micrographs. (A) Spines of fifth segment, lateral view; (B) magnified view of spines in (A), showing digitiform bosses on ventral row of spines; (C) dorsal view of individual with developing stolon 1; (D) dorsal view of paratomic chain with stock, stolon 2 (St2) and stolon 1 (St1); (E) lateral view of paratomic chain with stock, stolon 2, stolon 1 and developing secondary stolon (2‡St). Scale: 5 mm (A); 1 mm (B); 100 mm (C–E)." figureDoi="http://doi.org/10.5281/zenodo.5250252" httpUri="https://zenodo.org/record/5250252/files/figure.png" pageId="7" pageNumber="1345">2A, B</figureCitation>
).
</paragraph>
<paragraph id="DB6CCE6CFF97FD2654889AEE76D4FA10" blockId="7.[159,1116,989,1732]" pageId="7" pageNumber="1345">
Branchiae absent. Dorsal ciliary bands apparent from segment 3 continuing to terminal segments of stock individual (
<figureCitation id="43E8D2E9FF97FD2656289ACF76D8FA10" box="[615,754,1449,1473]" captionStart="FIG" captionStartId="5.[159,175,1364,1386]" captionTargetBox="[159,1116,171,1321]" captionTargetId="figure-51@5.[159,1116,170,1325]" captionTargetPageId="5" captionText="FIG. 2. Polydorella kamakamai new species: paratypes (USNM 1008712), SEM micrographs. (A) Spines of fifth segment, lateral view; (B) magnified view of spines in (A), showing digitiform bosses on ventral row of spines; (C) dorsal view of individual with developing stolon 1; (D) dorsal view of paratomic chain with stock, stolon 2 (St2) and stolon 1 (St1); (E) lateral view of paratomic chain with stock, stolon 2, stolon 1 and developing secondary stolon (2‡St). Scale: 5 mm (A); 1 mm (B); 100 mm (C–E)." figureDoi="http://doi.org/10.5281/zenodo.5250252" httpUri="https://zenodo.org/record/5250252/files/figure.png" pageId="7" pageNumber="1345">figure 2C–E</figureCitation>
).
</paragraph>
<paragraph id="DB6CCE6CFF97FD2654889AAF7769F9D3" blockId="7.[159,1116,989,1732]" pageId="7" pageNumber="1345">Narrow pharynx to segment 7 leading to broad intestine, narrowing in posterior segments; without gizzard-like structure in digestive tract.</paragraph>
<paragraph id="DB6CCE6CFF97FD265488996C76BAF992" blockId="7.[159,1116,989,1732]" pageId="7" pageNumber="1345">
Pygidium reduced, knob-shaped (
<figureCitation id="43E8D2E9FF97FD26561A996C7696F9F3" box="[597,700,1546,1570]" captionStart="FIG" captionStartId="3.[159,175,1286,1308]" captionTargetBox="[159,1115,172,1246]" captionTargetId="figure-98@3.[159,1117,170,1248]" captionTargetPageId="3" captionText="FIG. 1. Polydorella kamakamai new species: holotype (USNM 1008710) and paratypes (AMNH 4347). (A) Dorsal view of paratype exhibiting sexual reproduction (developing eggs in segments 13–15 outlined in dotted lines; three single eggs shown via shading); (B) dorsal view of holotype, arrowhead indicating growth zone following segment 10; (C) capillary neurosetae from segment 4; (D) spines of fifth segment, ventral view; (E) four acicular and two capillary neurosetae from segment 4; (F) hooded hook from segment 8. Scale: 100 mm (A, B); 10mm (C–F)." figureDoi="http://doi.org/10.5281/zenodo.5250250" httpUri="https://zenodo.org/record/5250250/files/figure.png" pageId="7" pageNumber="1345">figure 1B</figureCitation>
); with large cells (
<emphasis id="E9A7127EFF97FD2657EF996C7791F9F0" bold="true" box="[928,955,1546,1569]" italics="true" pageId="7" pageNumber="1345">y</emphasis>
<quantity id="1C2B6389FF97FD2657F4996C77C3F9F3" box="[955,1001,1546,1570]" metricMagnitude="1" metricUnit="m" metricValue="3.0" pageId="7" pageNumber="1345" unit="m" value="30.0">
30
<emphasis id="E9A7127EFF97FD265795996C77C3F9F3" bold="true" box="[986,1001,1546,1570]" italics="true" pageId="7" pageNumber="1345">m</emphasis>
</quantity>
m) within pygidium and terminal segments (
<figureCitation id="43E8D2E9FF97FD265662994D76A8F992" box="[557,642,1579,1603]" captionStart="FIG" captionStartId="6.[154,170,1594,1616]" captionTargetBox="[161,1109,211,1550]" captionTargetId="figure-1@6.[154,1112,207,1555]" captionTargetPageId="6" captionText="FIG. 3. Polydorella kamakamai new species: paratype (USNM 1008711). Paratomic chain with five developing individuals. (A) Dorsal view; (B) ventral view. Segment numbers of stock, stolon 2 and stolon 1 indicated to left of ventral view. P, pygidium. Scale: 250 mm." figureDoi="http://doi.org/10.5281/zenodo.5250254" httpUri="https://zenodo.org/record/5250254/files/figure.png" pageId="7" pageNumber="1345">figure 3</figureCitation>
).
</paragraph>
<paragraph id="DB6CCE6CFF97FD295488992D707CFB55" blockId="7.[159,1116,989,1732]" lastBlockId="8.[154,1110,1132,1156]" lastPageId="8" lastPageNumber="1346" pageId="7" pageNumber="1345">
<emphasis id="E9A7127EFF97FD265488992D756DF9B2" box="[199,327,1611,1635]" italics="true" pageId="7" pageNumber="1345">Variability.</emphasis>
All specimens examined exhibited black pigmentation on the palps, peristomium, prostomium, surrounding the caruncle, sides of anterior and posterior segments, and on the pygidium. Pigmentation was most prominent on the dorsal side but was also present on the ventral side of segments. A bifid prostomium and caruncle extending to middle of segment two was observed in
</paragraph>
<caption id="8FAC9EE4FF98FD2954D59C0177EFFBE6" ID-DOI="http://doi.org/10.5281/zenodo.5250258" ID-Zenodo-Dep="5250258" httpUri="https://zenodo.org/record/5250258/files/figure.png" pageId="8" pageNumber="1346" startId="8.[154,170,871,893]" targetBox="[157,1109,175,830]" targetPageId="8">
<paragraph id="DB6CCE6CFF98FD2954D59C0177EFFBE6" blockId="8.[154,1111,871,1079]" pageId="8" pageNumber="1346">
FIG. 5.
<taxonomicName id="1CD3B5EFFF98FD2954BB9C0E75C9FCAC" authority="Williams, 2004" authorityName="Williams" authorityYear="2004" box="[244,483,872,893]" class="Polychaeta" family="Spionidae" genus="Polydorella" kingdom="Animalia" order="Spionida" pageId="8" pageNumber="1346" phylum="Annelida" rank="species" species="kamakamai" status="sp. nov.">
<emphasis id="E9A7127EFF98FD2954BB9C0E75C9FCAC" box="[244,483,872,893]" italics="true" pageId="8" pageNumber="1346">Polydorella kamakamai</emphasis>
</taxonomicName>
<taxonomicNameLabel id="F294AF05FF98FD2955A29C017642FCAC" box="[493,616,871,893]" pageId="8" pageNumber="1346" rank="species">new species</taxonomicNameLabel>
. Semi-diagrammatic view of process of asexual reproduction via paratomy. (A) Single stock individual without growth zone (left palp shown for comparison with developing palp buds; palps removed from remaining figures); (B) development of stolon 1 (St1) growth zone; (C) development of stolon 2 (St2) growth zone; (D) development of first 10 segments of stolon 1 and stolon 3 (St3) growth zone; secondary stolon (2
<emphasis id="E9A7127EFF98FD2956779C8A766BFBD3" bold="true" box="[568,577,1004,1026]" pageId="8" pageNumber="1346">‡</emphasis>
St) indicated by arrowhead; (E) paratomic chain of five developing individuals just prior to paratomic division between stolon 2 and stolon 1; site of paratomic division indicated by arrow. Scale: 250
<emphasis id="E9A7127EFF98FD2957D29B447780FBE6" bold="true" box="[925,938,1058,1079]" italics="true" pageId="8" pageNumber="1346">m</emphasis>
m.
</paragraph>
</caption>
<paragraph id="DB6CCE6CFF98FD2954D59BE976B8FAFE" blockId="8.[154,1111,1167,1495]" pageId="8" pageNumber="1346">all specimens; no specimens possessed branchiae. Acicular neurosetae were found in approximately vertical rows reaching four in number on segment 5, accompanied by a dorsal row of typically four to six capillary neurosetae. The number of acicular neurosetae was reduced to two or three in segments 6 and 7, accompanied by one or two capillary neurosetae.</paragraph>
<paragraph id="DB6CCE6CFF98FD29548D9A51707CFA06" blockId="8.[154,1111,1167,1495]" pageId="8" pageNumber="1346">
The fifth segment contained four to seven major spines in both the dorsal and ventral rows; 20% of the specimens examined (
<emphasis id="E9A7127EFF98FD2956B39A3F773AFAA0" box="[764,784,1369,1393]" italics="true" pageId="8" pageNumber="1346">N</emphasis>
~50) possessed one more additional spine in the ventral row than the dorsal row; all other specimens possessed equivalent numbers of spines in both rows. Hooded hooks numbered three or four on segment 8, reaching a maximum of three to five hooded hooks on
</paragraph>
<paragraph id="DB6CCE6CFF98FD2954D59A847620F9EF" blockId="8.[154,1111,1506,1598]" pageId="8" pageNumber="1346">
segments 8–11. The pygidium was variable, from a reduced, knob-shape (
<figureCitation id="43E8D2E9FF98FD2957909A84707BFA2B" box="[991,1105,1506,1530]" captionStart="FIG" captionStartId="3.[159,175,1286,1308]" captionTargetBox="[159,1115,172,1246]" captionTargetId="figure-98@3.[159,1117,170,1248]" captionTargetPageId="3" captionText="FIG. 1. Polydorella kamakamai new species: holotype (USNM 1008710) and paratypes (AMNH 4347). (A) Dorsal view of paratype exhibiting sexual reproduction (developing eggs in segments 13–15 outlined in dotted lines; three single eggs shown via shading); (B) dorsal view of holotype, arrowhead indicating growth zone following segment 10; (C) capillary neurosetae from segment 4; (D) spines of fifth segment, ventral view; (E) four acicular and two capillary neurosetae from segment 4; (F) hooded hook from segment 8. Scale: 100 mm (A, B); 10mm (C–F)." figureDoi="http://doi.org/10.5281/zenodo.5250250" httpUri="https://zenodo.org/record/5250250/files/figure.png" pageId="8" pageNumber="1346">figures 1B</figureCitation>
,
<figureCitation id="43E8D2E9FF98FD2954D5996274C2F9CD" box="[154,232,1540,1564]" captionStart="FIG" captionStartId="5.[159,175,1364,1386]" captionTargetBox="[159,1116,171,1321]" captionTargetId="figure-51@5.[159,1116,170,1325]" captionTargetPageId="5" captionText="FIG. 2. Polydorella kamakamai new species: paratypes (USNM 1008712), SEM micrographs. (A) Spines of fifth segment, lateral view; (B) magnified view of spines in (A), showing digitiform bosses on ventral row of spines; (C) dorsal view of individual with developing stolon 1; (D) dorsal view of paratomic chain with stock, stolon 2 (St2) and stolon 1 (St1); (E) lateral view of paratomic chain with stock, stolon 2, stolon 1 and developing secondary stolon (2‡St). Scale: 5 mm (A); 1 mm (B); 100 mm (C–E)." figureDoi="http://doi.org/10.5281/zenodo.5250252" httpUri="https://zenodo.org/record/5250252/files/figure.png" pageId="8" pageNumber="1346">2C, D</figureCitation>
) to a more elongate conical shape (
<figureCitation id="43E8D2E9FF98FD2956F599627725F9CD" box="[698,783,1540,1564]" captionStart="FIG" captionStartId="6.[154,170,1594,1616]" captionTargetBox="[161,1109,211,1550]" captionTargetId="figure-1@6.[154,1112,207,1555]" captionTargetPageId="6" captionText="FIG. 3. Polydorella kamakamai new species: paratype (USNM 1008711). Paratomic chain with five developing individuals. (A) Dorsal view; (B) ventral view. Segment numbers of stock, stolon 2 and stolon 1 indicated to left of ventral view. P, pygidium. Scale: 250 mm." figureDoi="http://doi.org/10.5281/zenodo.5250254" httpUri="https://zenodo.org/record/5250254/files/figure.png" pageId="8" pageNumber="1346">figure 3</figureCitation>
); small bumps were often observed surrounding the anus.
</paragraph>
<paragraph id="DB6CCE6CFF98FD2854D599EB7528FF17" blockId="8.[154,552,1677,1732]" lastBlockId="9.[159,1116,174,326]" lastPageId="9" lastPageNumber="1347" pageId="8" pageNumber="1346">
<emphasis id="E9A7127EFF98FD2954D599EB750AF974" box="[154,288,1677,1701]" italics="true" pageId="8" pageNumber="1346">Distribution</emphasis>
<collectingCountry id="A3C48EFCFF98FD29548D99CA7569F915" box="[194,323,1708,1732]" name="Philippines" pageId="8" pageNumber="1346">Philippines</collectingCountry>
:
<collectingRegion id="1917008EFF98FD29551E99CA760EF915" box="[337,548,1708,1732]" country="Philippines" name="Mindoro Oriental" pageId="8" pageNumber="1346">Oriental Mindoro</collectingRegion>
.
<emphasis id="E9A7127EFF99FD2854D09FC87528FF17" box="[159,258,174,198]" italics="true" pageId="9" pageNumber="1347">Remarks</emphasis>
</paragraph>
<paragraph id="DB6CCE6CFF99FD2854889FA87707FE97" blockId="9.[159,1116,174,326]" pageId="9" pageNumber="1347">
<taxonomicName id="1CD3B5EFFF99FD2854889FA875FDFF37" authorityName="Williams" authorityYear="2004" box="[199,471,206,230]" class="Polychaeta" family="Spionidae" genus="Polydorella" kingdom="Animalia" order="Spionida" pageId="9" pageNumber="1347" phylum="Annelida" rank="species" species="kamakamai">
<emphasis id="E9A7127EFF99FD2854889FA875FDFF37" box="[199,471,206,230]" italics="true" pageId="9" pageNumber="1347">Polydorella kamakamai</emphasis>
</taxonomicName>
is unique within the genus in possession of acicular neurosetae in segments 2–7 and the fifth segment containing ventral row of major spines with digitiform bosses. The species is further differentiated from
<taxonomicName id="1CD3B5EFFF99FD2857869E68707FFEF7" authorityName="Radashevsky" authorityYear="1996" box="[969,1109,270,294]" class="Polychaeta" family="Spionidae" genus="Polydorella" kingdom="Animalia" order="Spionida" pageId="9" pageNumber="1347" phylum="Annelida" rank="species" species="dawydoffi">
<emphasis id="E9A7127EFF99FD2857869E68707FFEF7" box="[969,1109,270,294]" italics="true" pageId="9" pageNumber="1347">P. dawydoffi</emphasis>
</taxonomicName>
,
<taxonomicName id="1CD3B5EFFF99FD2854D09E487508FE97" authorityName="Augener" authorityYear="1914" box="[159,290,302,326]" class="Polychaeta" family="Spionidae" genus="Polydorella" kingdom="Animalia" order="Spionida" pageId="9" pageNumber="1347" phylum="Annelida" rank="species" species="prolifera">
<emphasis id="E9A7127EFF99FD2854D09E487508FE97" box="[159,290,302,326]" italics="true" pageId="9" pageNumber="1347">P. prolifera</emphasis>
</taxonomicName>
and
<taxonomicName id="1CD3B5EFFF99FD28552D9E4875D6FE97" baseAuthorityName="Blake and Kudenov" baseAuthorityYear="1978" box="[354,508,302,326]" class="Polychaeta" family="Spionidae" genus="Polydorella" kingdom="Animalia" order="Spionida" pageId="9" pageNumber="1347" phylum="Annelida" rank="species" species="stolonifera">
<emphasis id="E9A7127EFF99FD28552D9E4875D6FE97" box="[354,508,302,326]" italics="true" pageId="9" pageNumber="1347">P. stolonifera</emphasis>
</taxonomicName>
by the lack of branchiae.
</paragraph>
<paragraph id="DB6CCE6CFF99FD2854D09E1C74D3FE43" blockId="9.[159,1116,378,788]" box="[159,249,378,402]" pageId="9" pageNumber="1347">
<emphasis id="E9A7127EFF99FD2854D09E1C74D3FE43" box="[159,249,378,402]" italics="true" pageId="9" pageNumber="1347">Ecology</emphasis>
</paragraph>
<paragraph id="DB6CCE6CFF99FD2854889EFC76CBFCC5" blockId="9.[159,1116,378,788]" pageId="9" pageNumber="1347">
<taxonomicName id="1CD3B5EFFF99FD2854889EFC75FEFE63" authorityName="Williams" authorityYear="2004" box="[199,468,410,434]" class="Polychaeta" family="Spionidae" genus="Polydorella" kingdom="Animalia" order="Spionida" pageId="9" pageNumber="1347" phylum="Annelida" rank="species" species="kamakamai">
<emphasis id="E9A7127EFF99FD2854889EFC75FEFE63" box="[199,468,410,434]" italics="true" pageId="9" pageNumber="1347">Polydorella kamakamai</emphasis>
</taxonomicName>
was found to construct small mud tubes (up to
<quantity id="1C2B6389FF99FD28505A9EFF707EFE60" box="[1045,1108,409,433]" metricMagnitude="-3" metricUnit="m" metricValue="5.0" pageId="9" pageNumber="1347" unit="mm" value="5.0">5 mm</quantity>
) on the surface of the sponge,
<taxonomicName id="1CD3B5EFFF99FD2856409EDC77AAFE03" baseAuthorityName="Thiele" baseAuthorityYear="1903" box="[527,896,442,466]" class="Demospongiae" family="Microcionidae" genus="Clathria" kingdom="Animalia" order="Poecilosclerida" pageId="9" pageNumber="1347" phylum="Porifera" rank="species" species="cervicornis" subGenus="Thalysias">
<emphasis id="E9A7127EFF99FD2856409EDC77AAFE03" box="[527,896,442,466]" italics="true" pageId="9" pageNumber="1347">Clathria (Thalysias) cervicornis</emphasis>
</taxonomicName>
. The worms were found in fairly high abundance, with approximately 280 worms on a sponge consisting of eight main branches approximately
<quantity id="1C2B6389FF99FD2857429E9C774FFDC3" box="[781,869,506,530]" metricMagnitude="-1" metricUnit="m" metricValue="1.0" pageId="9" pageNumber="1347" unit="mm" value="100.0">100 mm</quantity>
long and
<quantity id="1C2B6389FF99FD2857B89E9C701EFDC3" box="[1015,1076,506,530]" metricMagnitude="-3" metricUnit="m" metricValue="7.0" pageId="9" pageNumber="1347" unit="mm" value="7.0">7 mm</quantity>
in diameter (
<emphasis id="E9A7127EFF99FD2855589D7D7518FDE3" bold="true" box="[279,306,539,562]" italics="true" pageId="9" pageNumber="1347">y</emphasis>
1892.0 cm
<superScript id="2CA66324FF99FD2855ED9D737586FDF4" attach="right" box="[418,428,533,549]" fontSize="7" pageId="9" pageNumber="1347">2</superScript>
). Some specimens contained material in the gut the same colour as the preserved sponge; additional behavioural and histological studies are required to determine if the worms feed on the tissue of sponges as has been documented in some other polychaetes (see
<bibRefCitation id="BF42B39DFF99FD2856F69D1D77D5FD42" author="MARTIN, D. &amp; BRITAYEV, T. A." box="[697,1023,635,659]" pageId="9" pageNumber="1347" pagination="217 - 340" refId="ref8921" refString="MARTIN, D. and BRITAYEV, T. A., 1998, Symbiotic polychaetes: Review of known species, Oceanography and Marine Biology. An Annual Review, 36, 217 - 340." type="journal article" year="1998">Martín and Britayev, 1998</bibRefCitation>
). Many tubes contained two individuals with dorsal sides apposed and anterior ends facing in opposite directions; tubes containing greater than two individuals were not observed. When disturbed, worms were able to move quickly out of their tubes and efficiently crawl across the surface of the sponge.
</paragraph>
<paragraph id="DB6CCE6CFF99FD2854D09C2E75BAFCB1" blockId="9.[159,1116,840,1732]" box="[159,400,840,864]" pageId="9" pageNumber="1347">
<emphasis id="E9A7127EFF99FD2854D09C2E75BAFCB1" box="[159,400,840,864]" italics="true" pageId="9" pageNumber="1347">Asexual reproduction</emphasis>
</paragraph>
<paragraph id="DB6CCE6CFF99FD2854889C01752FFB13" blockId="9.[159,1116,840,1732]" pageId="9" pageNumber="1347">
Most specimens examined (
<emphasis id="E9A7127EFF99FD2856469C0E7637FC51" box="[521,541,872,896]" italics="true" pageId="9" pageNumber="1347">N</emphasis>
~290) exhibited some stage of asexual reproduction via paratomy (
<figureCitation id="43E8D2E9FF99FD2855CA9CEE75F1FC71" box="[389,475,904,928]" captionStart="FIG" captionStartId="7.[159,175,875,897]" captionTargetBox="[309,969,173,834]" captionTargetId="figure-255@7.[306,970,170,837]" captionTargetPageId="7" captionText="FIG. 4. Polydorella kamakamai new species. Percentages of single individuals and paratomic chains of two, three, four and five or more individuals in colonies from the Philippines (N~290)." figureDoi="http://doi.org/10.5281/zenodo.5250256" httpUri="https://zenodo.org/record/5250256/files/figure.png" pageId="9" pageNumber="1347">figure 4</figureCitation>
). Single parental or stock individuals consist of 13–15 segments (
<figureCitation id="43E8D2E9FF99FD2855579CCE75AFFC11" box="[280,389,936,960]" captionStart="FIG" captionStartId="8.[154,170,871,893]" captionTargetBox="[157,1109,175,830]" captionTargetId="figure-173@8.[154,1112,170,833]" captionTargetPageId="8" captionText="FIG. 5. Polydorella kamakamai new species. Semi-diagrammatic view of process of asexual reproduction via paratomy. (A) Single stock individual without growth zone (left palp shown for comparison with developing palp buds; palps removed from remaining figures); (B) development of stolon 1 (St1) growth zone; (C) development of stolon 2 (St2) growth zone; (D) development of first 10 segments of stolon 1 and stolon 3 (St3) growth zone; secondary stolon (2‡St) indicated by arrowhead; (E) paratomic chain of five developing individuals just prior to paratomic division between stolon 2 and stolon 1; site of paratomic division indicated by arrow. Scale: 250 mm." figureDoi="http://doi.org/10.5281/zenodo.5250258" httpUri="https://zenodo.org/record/5250258/files/figure.png" pageId="9" pageNumber="1347">figure 5A</figureCitation>
). Stock individuals then exhibit a growth zone which follows segment 10 (
<figureCitation id="43E8D2E9FF99FD2855779CAE7583FC31" box="[312,425,968,992]" captionStart="FIG" captionStartId="3.[159,175,1286,1308]" captionTargetBox="[159,1115,172,1246]" captionTargetId="figure-98@3.[159,1117,170,1248]" captionTargetPageId="3" captionText="FIG. 1. Polydorella kamakamai new species: holotype (USNM 1008710) and paratypes (AMNH 4347). (A) Dorsal view of paratype exhibiting sexual reproduction (developing eggs in segments 13–15 outlined in dotted lines; three single eggs shown via shading); (B) dorsal view of holotype, arrowhead indicating growth zone following segment 10; (C) capillary neurosetae from segment 4; (D) spines of fifth segment, ventral view; (E) four acicular and two capillary neurosetae from segment 4; (F) hooded hook from segment 8. Scale: 100 mm (A, B); 10mm (C–F)." figureDoi="http://doi.org/10.5281/zenodo.5250250" httpUri="https://zenodo.org/record/5250250/files/figure.png" pageId="9" pageNumber="1347">figures 1B</figureCitation>
,
<figureCitation id="43E8D2E9FF99FD2855F39CAE75F5FC31" box="[444,479,968,992]" captionStart="FIG" captionStartId="5.[159,175,1364,1386]" captionTargetBox="[159,1116,171,1321]" captionTargetId="figure-51@5.[159,1116,170,1325]" captionTargetPageId="5" captionText="FIG. 2. Polydorella kamakamai new species: paratypes (USNM 1008712), SEM micrographs. (A) Spines of fifth segment, lateral view; (B) magnified view of spines in (A), showing digitiform bosses on ventral row of spines; (C) dorsal view of individual with developing stolon 1; (D) dorsal view of paratomic chain with stock, stolon 2 (St2) and stolon 1 (St1); (E) lateral view of paratomic chain with stock, stolon 2, stolon 1 and developing secondary stolon (2‡St). Scale: 5 mm (A); 1 mm (B); 100 mm (C–E)." figureDoi="http://doi.org/10.5281/zenodo.5250252" httpUri="https://zenodo.org/record/5250252/files/figure.png" pageId="9" pageNumber="1347">2C</figureCitation>
,
<figureCitation id="43E8D2E9FF99FD2855BF9CAE7639FC31" box="[496,531,968,992]" captionStart="FIG" captionStartId="8.[154,170,871,893]" captionTargetBox="[157,1109,175,830]" captionTargetId="figure-173@8.[154,1112,170,833]" captionTargetPageId="8" captionText="FIG. 5. Polydorella kamakamai new species. Semi-diagrammatic view of process of asexual reproduction via paratomy. (A) Single stock individual without growth zone (left palp shown for comparison with developing palp buds; palps removed from remaining figures); (B) development of stolon 1 (St1) growth zone; (C) development of stolon 2 (St2) growth zone; (D) development of first 10 segments of stolon 1 and stolon 3 (St3) growth zone; secondary stolon (2‡St) indicated by arrowhead; (E) paratomic chain of five developing individuals just prior to paratomic division between stolon 2 and stolon 1; site of paratomic division indicated by arrow. Scale: 250 mm." figureDoi="http://doi.org/10.5281/zenodo.5250258" httpUri="https://zenodo.org/record/5250258/files/figure.png" pageId="9" pageNumber="1347">5B</figureCitation>
). The growth zone is distinct from anterior and posterior segments in its opaque colouring and smooth form. The opaque colouring of the growth zone may come from the breakdown of muscular material as the zone prepares for the formation of a new stolon body (
<bibRefCitation id="BF42B39DFF99FD2856A49B4F707AFB90" author="SCHROEDER, P. C. &amp; HERMANS, C. O." box="[747,1104,1065,1089]" pageId="9" pageNumber="1347" refId="ref9251" refString="SCHROEDER, P. C. and HERMANS, C. O., 1975, Annelida: Polychaeta, in A. C. Giese and J. S. Pearse (eds) Reproduction of Marine Invertebrates (New York: Academic Press)." type="book" year="1975">Schroeder and Hermans, 1975</bibRefCitation>
). The growth zone exhibits two sections: an anterior section that will become the new segment 11 (denoted as 11
<emphasis id="E9A7127EFF99FD28559C9B0E75F3FB50" box="[467,473,1128,1153]" italics="true" pageId="9" pageNumber="1347">’</emphasis>
, following
<bibRefCitation id="BF42B39DFF99FD2856109B0F776CFB50" author="RADASHEVSKY, V. I." box="[607,838,1129,1153]" pageId="9" pageNumber="1347" pagination="684 - 693" refId="ref9088" refString="RADASHEVSKY, V. I., 1996, Morphology, ecology and asexual reproduction of a new Polydorella species (Polychaeta; Spionidae) from the South China Sea, Bulletin of Marine Science, 58, 684 - 693." type="journal article" year="1996">Radashevsky (1996)</bibRefCitation>
where
<emphasis id="E9A7127EFF99FD2857D29B0E7789FB50" box="[925,931,1128,1153]" italics="true" pageId="9" pageNumber="1347">’</emphasis>
indicates newly developed stock segments) and the posterior portion which produces the developing stolon 1.
</paragraph>
<paragraph id="DB6CCE6CFF99FD2854889BAC7602F955" blockId="9.[159,1116,840,1732]" pageId="9" pageNumber="1347">
After further development of the growth zone, palp buds are formed and the new stock segment anterior to the head of the developing stolon 1 exhibits setae. New segments of the stolon body then appear to form simultaneously (overt distinction of 10 segments displayed at once from subdivision of the elongated growth zone). Ultimately, stolon 1 consists of 10 new anterior segments and three to four posterior segments derived from the stock individual (
<figureCitation id="43E8D2E9FF99FD28572F9A0D77E7FA52" box="[864,973,1387,1411]" captionStart="FIG" captionStartId="8.[154,170,871,893]" captionTargetBox="[157,1109,175,830]" captionTargetId="figure-173@8.[154,1112,170,833]" captionTargetPageId="8" captionText="FIG. 5. Polydorella kamakamai new species. Semi-diagrammatic view of process of asexual reproduction via paratomy. (A) Single stock individual without growth zone (left palp shown for comparison with developing palp buds; palps removed from remaining figures); (B) development of stolon 1 (St1) growth zone; (C) development of stolon 2 (St2) growth zone; (D) development of first 10 segments of stolon 1 and stolon 3 (St3) growth zone; secondary stolon (2‡St) indicated by arrowhead; (E) paratomic chain of five developing individuals just prior to paratomic division between stolon 2 and stolon 1; site of paratomic division indicated by arrow. Scale: 250 mm." figureDoi="http://doi.org/10.5281/zenodo.5250258" httpUri="https://zenodo.org/record/5250258/files/figure.png" pageId="9" pageNumber="1347">figure 5D</figureCitation>
). Stolon 2 is formed following 11
<emphasis id="E9A7127EFF99FD2855C09AEC75BFFA72" box="[399,405,1418,1443]" italics="true" pageId="9" pageNumber="1347">’</emphasis>
. Prior to complete development of the first 10 segments of stolon 1 (i.e. lacking externally observable setae), a secondary growth zone is formed following segment 10. Formation of stolon 2 proceeds as described above and the secondary growth zone of stolon 1 then exhibits palp buds (
<figureCitation id="43E8D2E9FF99FD2857F19A8D707DF9D2" box="[958,1111,1515,1539]" captionStart="FIG" captionStartId="5.[159,175,1364,1386]" captionTargetBox="[159,1116,171,1321]" captionTargetId="figure-51@5.[159,1116,170,1325]" captionTargetPageId="5" captionText="FIG. 2. Polydorella kamakamai new species: paratypes (USNM 1008712), SEM micrographs. (A) Spines of fifth segment, lateral view; (B) magnified view of spines in (A), showing digitiform bosses on ventral row of spines; (C) dorsal view of individual with developing stolon 1; (D) dorsal view of paratomic chain with stock, stolon 2 (St2) and stolon 1 (St1); (E) lateral view of paratomic chain with stock, stolon 2, stolon 1 and developing secondary stolon (2‡St). Scale: 5 mm (A); 1 mm (B); 100 mm (C–E)." figureDoi="http://doi.org/10.5281/zenodo.5250252" httpUri="https://zenodo.org/record/5250252/files/figure.png" pageId="9" pageNumber="1347">figures 2D, E</figureCitation>
,
<figureCitation id="43E8D2E9FF99FD2854D0996A74ECF9F5" box="[159,198,1548,1572]" captionStart="FIG" captionStartId="8.[154,170,871,893]" captionTargetBox="[157,1109,175,830]" captionTargetId="figure-173@8.[154,1112,170,833]" captionTargetPageId="8" captionText="FIG. 5. Polydorella kamakamai new species. Semi-diagrammatic view of process of asexual reproduction via paratomy. (A) Single stock individual without growth zone (left palp shown for comparison with developing palp buds; palps removed from remaining figures); (B) development of stolon 1 (St1) growth zone; (C) development of stolon 2 (St2) growth zone; (D) development of first 10 segments of stolon 1 and stolon 3 (St3) growth zone; secondary stolon (2‡St) indicated by arrowhead; (E) paratomic chain of five developing individuals just prior to paratomic division between stolon 2 and stolon 1; site of paratomic division indicated by arrow. Scale: 250 mm." figureDoi="http://doi.org/10.5281/zenodo.5250258" httpUri="https://zenodo.org/record/5250258/files/figure.png" pageId="9" pageNumber="1347">5D</figureCitation>
). A third stolon (stolon 3) is observed in some individuals behind 11
<emphasis id="E9A7127EFF99FD2857B8996D77D7F9F5" box="[1015,1021,1547,1572]" italics="true" pageId="9" pageNumber="1347">’</emphasis>
. In this way, a chain of five developing individuals is formed (stock, stolon 3, stolon 2, stolon 1 and the secondary stolon) (
<figureCitation id="43E8D2E9FF99FD285613992A76D2F9B5" box="[604,760,1612,1636]" captionStart="FIG" captionStartId="6.[154,170,1594,1616]" captionTargetBox="[161,1109,211,1550]" captionTargetId="figure-1@6.[154,1112,207,1555]" captionTargetPageId="6" captionText="FIG. 3. Polydorella kamakamai new species: paratype (USNM 1008711). Paratomic chain with five developing individuals. (A) Dorsal view; (B) ventral view. Segment numbers of stock, stolon 2 and stolon 1 indicated to left of ventral view. P, pygidium. Scale: 250 mm." figureDoi="http://doi.org/10.5281/zenodo.5250254" httpUri="https://zenodo.org/record/5250254/files/figure.png" pageId="9" pageNumber="1347">figures 3A, B</figureCitation>
,
<figureCitation id="43E8D2E9FF99FD285744992A7707F9B5" box="[779,813,1612,1636]" captionStart="FIG" captionStartId="8.[154,170,871,893]" captionTargetBox="[157,1109,175,830]" captionTargetId="figure-173@8.[154,1112,170,833]" captionTargetPageId="8" captionText="FIG. 5. Polydorella kamakamai new species. Semi-diagrammatic view of process of asexual reproduction via paratomy. (A) Single stock individual without growth zone (left palp shown for comparison with developing palp buds; palps removed from remaining figures); (B) development of stolon 1 (St1) growth zone; (C) development of stolon 2 (St2) growth zone; (D) development of first 10 segments of stolon 1 and stolon 3 (St3) growth zone; secondary stolon (2‡St) indicated by arrowhead; (E) paratomic chain of five developing individuals just prior to paratomic division between stolon 2 and stolon 1; site of paratomic division indicated by arrow. Scale: 250 mm." figureDoi="http://doi.org/10.5281/zenodo.5250258" httpUri="https://zenodo.org/record/5250258/files/figure.png" pageId="9" pageNumber="1347">5E</figureCitation>
). The process of asexual reproduction is shown in
<figureCitation id="43E8D2E9FF99FD285580990A760EF955" box="[463,548,1644,1668]" captionStart="FIG" captionStartId="8.[154,170,871,893]" captionTargetBox="[157,1109,175,830]" captionTargetId="figure-173@8.[154,1112,170,833]" captionTargetPageId="8" captionText="FIG. 5. Polydorella kamakamai new species. Semi-diagrammatic view of process of asexual reproduction via paratomy. (A) Single stock individual without growth zone (left palp shown for comparison with developing palp buds; palps removed from remaining figures); (B) development of stolon 1 (St1) growth zone; (C) development of stolon 2 (St2) growth zone; (D) development of first 10 segments of stolon 1 and stolon 3 (St3) growth zone; secondary stolon (2‡St) indicated by arrowhead; (E) paratomic chain of five developing individuals just prior to paratomic division between stolon 2 and stolon 1; site of paratomic division indicated by arrow. Scale: 250 mm." figureDoi="http://doi.org/10.5281/zenodo.5250258" httpUri="https://zenodo.org/record/5250258/files/figure.png" pageId="9" pageNumber="1347">figure 5</figureCitation>
.
</paragraph>
<paragraph id="DB6CCE6CFF99FD2B548899EA752FFDAC" blockId="9.[159,1116,840,1732]" lastBlockId="10.[154,1111,173,804]" lastPageId="10" lastPageNumber="1348" pageId="9" pageNumber="1347">
It appears that in most cases paratomic division takes place when four or five individuals are present in a chain. Stolon 1 (with secondary stolon) breaks from the stock individual (stolon 3 and stolon 2) (
<figureCitation id="43E8D2E9FF9AFD2B563A9FCB76F4FF14" box="[629,734,173,197]" captionStart="FIG" captionStartId="8.[154,170,871,893]" captionTargetBox="[157,1109,175,830]" captionTargetId="figure-173@8.[154,1112,170,833]" captionTargetPageId="8" captionText="FIG. 5. Polydorella kamakamai new species. Semi-diagrammatic view of process of asexual reproduction via paratomy. (A) Single stock individual without growth zone (left palp shown for comparison with developing palp buds; palps removed from remaining figures); (B) development of stolon 1 (St1) growth zone; (C) development of stolon 2 (St2) growth zone; (D) development of first 10 segments of stolon 1 and stolon 3 (St3) growth zone; secondary stolon (2‡St) indicated by arrowhead; (E) paratomic chain of five developing individuals just prior to paratomic division between stolon 2 and stolon 1; site of paratomic division indicated by arrow. Scale: 250 mm." figureDoi="http://doi.org/10.5281/zenodo.5250258" httpUri="https://zenodo.org/record/5250258/files/figure.png" pageId="10" pageNumber="1348">figure 5E</figureCitation>
). Prior to division, the posterior segments of stolon 2 are found in a wedge-shape under the head of stolon 1 (
<figureCitation id="43E8D2E9FF9AFD2B54EE9F97753EFED8" box="[161,276,241,265]" captionStart="FIG" captionStartId="5.[159,175,1364,1386]" captionTargetBox="[159,1116,171,1321]" captionTargetId="figure-51@5.[159,1116,170,1325]" captionTargetPageId="5" captionText="FIG. 2. Polydorella kamakamai new species: paratypes (USNM 1008712), SEM micrographs. (A) Spines of fifth segment, lateral view; (B) magnified view of spines in (A), showing digitiform bosses on ventral row of spines; (C) dorsal view of individual with developing stolon 1; (D) dorsal view of paratomic chain with stock, stolon 2 (St2) and stolon 1 (St1); (E) lateral view of paratomic chain with stock, stolon 2, stolon 1 and developing secondary stolon (2‡St). Scale: 5 mm (A); 1 mm (B); 100 mm (C–E)." figureDoi="http://doi.org/10.5281/zenodo.5250252" httpUri="https://zenodo.org/record/5250252/files/figure.png" pageId="10" pageNumber="1348">figures 2E</figureCitation>
,
<figureCitation id="43E8D2E9FF9AFD2B556F9F977568FED8" box="[288,322,241,265]" captionStart="FIG" captionStartId="6.[154,170,1594,1616]" captionTargetBox="[161,1109,211,1550]" captionTargetId="figure-1@6.[154,1112,207,1555]" captionTargetPageId="6" captionText="FIG. 3. Polydorella kamakamai new species: paratype (USNM 1008711). Paratomic chain with five developing individuals. (A) Dorsal view; (B) ventral view. Segment numbers of stock, stolon 2 and stolon 1 indicated to left of ventral view. P, pygidium. Scale: 250 mm." figureDoi="http://doi.org/10.5281/zenodo.5250254" httpUri="https://zenodo.org/record/5250254/files/figure.png" pageId="10" pageNumber="1348">3B</figureCitation>
). At this time, palps of stolon 1 are approximately one-half to threequarters the length of stock individuals. Following division, the pygidium is fully formed on the posterior end of stolon 2. Stolon 1 (with secondary stolon) and stock (with stolon 3 and stolon 2) individuals then repeat the course of asexual reproduction described above. Following division, the worms apparently do not immediately leave their original tubes and construct new ones, as evidenced by tubes containing two individuals. The division of paratomic chains with stock individuals lacking a stolon 3 or stolon 2 leads to the presence of single individuals within the population. In only seven (2.4%) cases were the stock individuals within paratomic chains observed to possess 13–15 stock segments prior to division; this low number explains the relative rarity of single individuals lacking growth zones (
<figureCitation id="43E8D2E9FF9AFD2B54EE9D0374D2FDAC" box="[161,248,613,637]" captionStart="FIG" captionStartId="7.[159,175,875,897]" captionTargetBox="[309,969,173,834]" captionTargetId="figure-255@7.[306,970,170,837]" captionTargetPageId="7" captionText="FIG. 4. Polydorella kamakamai new species. Percentages of single individuals and paratomic chains of two, three, four and five or more individuals in colonies from the Philippines (N~290)." figureDoi="http://doi.org/10.5281/zenodo.5250256" httpUri="https://zenodo.org/record/5250256/files/figure.png" pageId="10" pageNumber="1348">figure 4</figureCitation>
).
</paragraph>
<paragraph id="DB6CCE6CFF9AFD2B548D9DE375E5FCF5" blockId="10.[154,1111,173,804]" pageId="10" pageNumber="1348">
The colonies were dominated by specimens composed of chains of two individuals (sometimes with developing growth zones); chains of three and four individuals were approximately half as abundant as chains of two individuals (
<figureCitation id="43E8D2E9FF9AFD2B54EE9D8C74D2FCD3" box="[161,248,746,770]" captionStart="FIG" captionStartId="7.[159,175,875,897]" captionTargetBox="[309,969,173,834]" captionTargetId="figure-255@7.[306,970,170,837]" captionTargetPageId="7" captionText="FIG. 4. Polydorella kamakamai new species. Percentages of single individuals and paratomic chains of two, three, four and five or more individuals in colonies from the Philippines (N~290)." figureDoi="http://doi.org/10.5281/zenodo.5250256" httpUri="https://zenodo.org/record/5250256/files/figure.png" pageId="10" pageNumber="1348">figure 4</figureCitation>
). Only rarely were single individuals and chains of five individuals or greater observed (
<figureCitation id="43E8D2E9FF9AFD2B55249C6A75E8FCF5" box="[363,450,780,804]" captionStart="FIG" captionStartId="7.[159,175,875,897]" captionTargetBox="[309,969,173,834]" captionTargetId="figure-255@7.[306,970,170,837]" captionTargetPageId="7" captionText="FIG. 4. Polydorella kamakamai new species. Percentages of single individuals and paratomic chains of two, three, four and five or more individuals in colonies from the Philippines (N~290)." figureDoi="http://doi.org/10.5281/zenodo.5250256" httpUri="https://zenodo.org/record/5250256/files/figure.png" pageId="10" pageNumber="1348">figure 4</figureCitation>
).
</paragraph>
<paragraph id="DB6CCE6CFF9AFD2B54D59C3E7557FCA1" blockId="10.[154,1111,856,1080]" box="[154,381,856,880]" pageId="10" pageNumber="1348">
<emphasis id="E9A7127EFF9AFD2B54D59C3E7557FCA1" box="[154,381,856,880]" italics="true" pageId="10" pageNumber="1348">Sexual reproduction</emphasis>
</paragraph>
<paragraph id="DB6CCE6CFF9AFD2B548D9C11707DFBE9" blockId="10.[154,1111,856,1080]" pageId="10" pageNumber="1348">
In addition to asexual reproduction,
<specimenCount id="CDD505E5FF9AFD2B56319C117731FC5E" box="[638,795,887,911]" pageId="10" pageNumber="1348" type="generic">one specimen</specimenCount>
composed of 15 segments was found with eggs (
<figureCitation id="43E8D2E9FF9AFD2B55EF9CFF7621FC60" box="[416,523,921,945]" captionStart="FIG" captionStartId="3.[159,175,1286,1308]" captionTargetBox="[159,1115,172,1246]" captionTargetId="figure-98@3.[159,1117,170,1248]" captionTargetPageId="3" captionText="FIG. 1. Polydorella kamakamai new species: holotype (USNM 1008710) and paratypes (AMNH 4347). (A) Dorsal view of paratype exhibiting sexual reproduction (developing eggs in segments 13–15 outlined in dotted lines; three single eggs shown via shading); (B) dorsal view of holotype, arrowhead indicating growth zone following segment 10; (C) capillary neurosetae from segment 4; (D) spines of fifth segment, ventral view; (E) four acicular and two capillary neurosetae from segment 4; (F) hooded hook from segment 8. Scale: 100 mm (A, B); 10mm (C–F)." figureDoi="http://doi.org/10.5281/zenodo.5250250" httpUri="https://zenodo.org/record/5250250/files/figure.png" pageId="10" pageNumber="1348">figure 1A</figureCitation>
). Segments 10–12 of the specimen were narrower than preceding segments (no growth zone was exhibited), while segments 13–15 were expanded and contained yolky eggs. Eggs were oval in shape; 43.3
<emphasis id="E9A7127EFF9AFD2B57949CBB77C0FC24" bold="true" box="[987,1002,989,1013]" italics="true" pageId="10" pageNumber="1348">m</emphasis>
m (¡5.8) by 62.2
<emphasis id="E9A7127EFF9AFD2B54BD9C99752BFBC6" bold="true" box="[242,257,1023,1047]" italics="true" pageId="10" pageNumber="1348">m</emphasis>
m (¡6.8) (
<emphasis id="E9A7127EFF9AFD2B55CF9C9975BEFBC6" box="[384,404,1023,1047]" italics="true" pageId="10" pageNumber="1348">N</emphasis>
~11). Based on the volume of the eggs within this specimen, segments 13–15 are estimated to contain 36–42, 26–34 and 15–
<specimenCount id="CDD505E5FF9AFD2B57289B467795FBE9" box="[871,959,1056,1080]" pageId="10" pageNumber="1348" type="egg">20 eggs</specimenCount>
, respectively.
</paragraph>
</subSubSection>
</treatment>
</document>
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