313 lines
47 KiB
XML
313 lines
47 KiB
XML
<document id="271A9A06A82200D370AA142FB042D50F" ID-DOI="10.11646/zootaxa.5453.2.1" ID-ISSN="1175-5326" ID-Zenodo-Dep="11233692" ID-ZooBank="B4856FA9-DE2F-4174-8F65-DB927813E48D" IM.bibliography_approvedBy="diego" IM.illustrations_approvedBy="diego" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="GgImagineBatch" IM.taxonomicNames_approvedBy="diego" IM.treatments_approvedBy="diego" checkinTime="1716282410815" checkinUser="plazi" docAuthor="Sawyer, Roy T." docDate="2024" docId="03BB0601BD0EFFFEFF1CB287FC08FA5C" docLanguage="en" docName="zootaxa.5453.2.1.pdf" docOrigin="Zootaxa 5453 (2)" docSource="http://dx.doi.org/10.11646/zootaxa.5453.2.1" docStyle="DocumentStyle:647186512141C8FC8976D5BCC54AEB7D.9:Zootaxa.2013-.journal_article" docStyleId="647186512141C8FC8976D5BCC54AEB7D" docStyleName="Zootaxa.2013-.journal_article" docStyleVersion="9" docTitle="Helobdella lineata Albemarle" docType="treatment" docVersion="3" lastPageNumber="164" masterDocId="FF827E79BD05FFF3FF8BB10DFFA5FFF4" masterDocTitle="Biology, morphology and taxonomy of a snail-feeding leech from North Carolina, USA, provisionally identified as Helobdella lineata (Verrill, 1874) (Glossiphoniidae): First evidence for extra-oral digestion in the Hirudinea" masterLastPageNumber="182" masterPageNumber="151" pageNumber="162" updateTime="1717163040250" updateUser="diego" zenodo-license-document="CLOSED">
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<mods:title id="E8E9A78B592680704082A9741BE5C749">Biology, morphology and taxonomy of a snail-feeding leech from North Carolina, USA, provisionally identified as Helobdella lineata (Verrill, 1874) (Glossiphoniidae): First evidence for extra-oral digestion in the Hirudinea</mods:title>
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<mods:namePart id="F55993902BFD540D446274A2D5A020E3">Sawyer, Roy T.</mods:namePart>
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<mods:title id="4ABDE1DD91768254D9740CF9F5DDC7D0">Zootaxa</mods:title>
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<mods:part id="A5AB655994D65CE859A73156B7246043">
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<mods:date id="DC0A5792126B24CF0470FFADDCC26066">2024</mods:date>
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<mods:number id="ADC2492ABB7ECE3079AD6839A5B6B7CB">2024-05-20</mods:number>
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<mods:url id="C90FD1E1749BC3F84693D89E354F3270">http://dx.doi.org/10.11646/zootaxa.5453.2.1</mods:url>
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<mods:identifier id="321BBB0050D2F58D7142B10FAC763CC7" type="DOI">10.11646/zootaxa.5453.2.1</mods:identifier>
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<mods:identifier id="BE3B7985ABD543C5425CD71CC25011E4" type="ISSN">1175-5326</mods:identifier>
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<treatment id="03BB0601BD0EFFFEFF1CB287FC08FA5C" LSID="urn:lsid:plazi:treatment:03BB0601BD0EFFFEFF1CB287FC08FA5C" httpUri="http://treatment.plazi.org/id/03BB0601BD0EFFFEFF1CB287FC08FA5C" lastPageId="13" lastPageNumber="164" pageId="11" pageNumber="162">
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<subSubSection id="C308E49CBD0EFFF8FF1CB287FD8BFC51" box="[151,558,906,933]" pageId="11" pageNumber="162" type="nomenclature">
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<paragraph id="8BADB717BD0EFFF8FF1CB287FD8BFC51" blockId="11.[151,558,906,933]" box="[151,558,906,933]" pageId="11" pageNumber="162">
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<heading id="D0E5007BBD0EFFF8FF1CB287FD8BFC51" bold="true" box="[151,558,906,933]" fontSize="11" level="1" pageId="11" pageNumber="162" reason="1">
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<emphasis id="B9666B05BD0EFFF8FF1CB287FD8BFC51" bold="true" box="[151,558,906,933]" pageId="11" pageNumber="162">
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Description of
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<taxonomicName id="4C12CC94BD0EFFF8FEC4B287FD8BFC51" authority="(Verrill, 1874) (Albemarle)" authorityName="Albemarle" baseAuthorityName="Verrill" baseAuthorityYear="1874" box="[335,558,906,933]" class="Clitellata" family="Glossiphoniidae" genus="Helobdella" kingdom="Animalia" order="Rhynchobdellida" pageId="11" pageNumber="162" phylum="Annelida" rank="species" species="lineata">
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<emphasis id="B9666B05BD0EFFF8FEC4B287FD8BFC51" bold="true" box="[335,558,906,933]" italics="true" pageId="11" pageNumber="162">Helobdella lineata</emphasis>
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</taxonomicName>
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</emphasis>
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</heading>
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</paragraph>
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</subSubSection>
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<subSubSection id="C308E49CBD0EFFFEFF1CB2DEFC08FA5C" lastPageId="13" lastPageNumber="164" pageId="11" pageNumber="162" type="description">
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<paragraph id="8BADB717BD0EFFF8FF1CB2DEFED5FC18" blockId="11.[151,368,978,1005]" box="[151,368,978,1005]" pageId="11" pageNumber="162">
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<heading id="D0E5007BBD0EFFF8FF1CB2DEFED5FC18" bold="true" box="[151,368,978,1005]" fontSize="11" level="1" pageId="11" pageNumber="162" reason="1">
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<emphasis id="B9666B05BD0EFFF8FF1CB2DEFED5FC18" bold="true" box="[151,368,978,1005]" pageId="11" pageNumber="162">External features</emphasis>
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</heading>
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</paragraph>
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<paragraph id="8BADB717BD0EFFF8FF1CB516FBF9FB1D" blockId="11.[151,1437,1050,2013]" pageId="11" pageNumber="162">
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<materialsCitation id="3B7ABD4ABD0EFFF8FF1CB516FBF2FB1D" county="Pasquotank County" location="Albemarle region" municipality="Albemarle region" pageId="11" pageNumber="162" specimenCount="392">
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A single adult specimen (HL-5A) is selected as being representative of the snail leech indigenous to the
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<location id="8ECDE1CCBD0EFFF8FAADB516FF7AFBAD" LSID="urn:lsid:plazi:treatment:03BB0601BD0EFFFEFF1CB287FC08FA5C:8ECDE1CCBD0EFFF8FAADB516FF7AFBAD" county="Pasquotank County" municipality="Albemarle region" name="Albemarle region" pageId="11" pageNumber="162">Albemarle region</location>
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(
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<figureCitation id="1329AB92BD0EFFF8FF78B532FE98FBAD" box="[243,317,1087,1113]" captionStart="FIGURE 4" captionStartId="5.[151,250,1904,1928]" captionTargetBox="[299,1288,774,1878]" captionTargetId="figure-275@5.[299,1288,774,1878]" captionTargetPageId="5" captionText="FIGURE 4. The archetype or reference specimen (HL-5A) on which the description of H. lineata is based. This specimen has faded and become translucent following preservation with 5% formalin. The type locality is in Pasquotank County, NC [36.191176, -76.228516] A, dorsal view. B, ventral view. Abbreviations: a, atrium; cs, caudal sucker; ic, intestinal caecum; pc, post caecum; pr, proboscis; r, rectum; rb, rectal bladder; sv, seminal vesicle; tp, tip of proboscis. Scale bar: 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.11233700" httpUri="https://zenodo.org/record/11233700/files/figure.png" pageId="11" pageNumber="162">Fig. 4</figureCitation>
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).
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<location id="8ECDE1CCBD0EFFF8FEDEB532FE24FBAD" LSID="urn:lsid:plazi:treatment:03BB0601BD0EFFFEFF1CB287FC08FA5C:8ECDE1CCBD0EFFF8FEDEB532FE24FBAD" box="[341,385,1087,1113]" county="Pasquotank County" municipality="Albemarle region" name="The" pageId="11" pageNumber="162">The</location>
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following description of this relatively large individual (
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<quantity id="4CEA1AF2BD0EFFF8FBA9B532FB28FBAD" box="[1058,1165,1087,1113]" metricMagnitude="-2" metricUnit="m" metricValue="1.28" pageId="11" pageNumber="162" unit="mm" value="12.8">12.8 mm</quantity>
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) (representing pigment variant #1) is supplemented by observations on eleven additional specimens collected at the same location (HL-5) in
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<collectingCounty id="62CCCF9BBD0EFFF8FF5EB58AFE16FB55" box="[213,435,1159,1185]" pageId="11" pageNumber="162">Pasquotank County</collectingCounty>
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. These twelve cohorts are remarkably homogeneous in size (
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<quantity id="4CEA1AF2BD0EFFF8FBE3B58AFAA1FB55" box="[1128,1284,1159,1185]" metricMagnitude="-2" metricUnit="m" metricValue="1.13" metricValueMax="1.28" metricValueMin="0.98" pageId="11" pageNumber="162" unit="mm" value="11.3" valueMax="12.8" valueMin="9.8">9.8–12.8 mm</quantity>
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), and state of feeding (empty gut) and reproduction (not gravid).
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<location id="8ECDE1CCBD0EFFF8FD40B5A6FC8FFB31" LSID="urn:lsid:plazi:treatment:03BB0601BD0EFFFEFF1CB287FC08FA5C:8ECDE1CCBD0EFFF8FD40B5A6FC8FFB31" box="[715,810,1195,1221]" county="Pasquotank County" municipality="Albemarle region" name="External" pageId="11" pageNumber="162">External</location>
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variability is based on examination of the additional
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<specimenCount id="9D147C9EBD0EFFF8FAF9B5A6FEA9FB1D" count="392" pageId="11" pageNumber="162" type="generic">392 specimens</specimenCount>
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of this species collected throughout the
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<collectingMunicipality id="6BC92D6DBD0EFFF8FD45B5C2FC36FB1D" box="[718,915,1231,1257]" pageId="11" pageNumber="162">Albemarle region</collectingMunicipality>
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during this study
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</materialsCitation>
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.
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</paragraph>
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<paragraph id="8BADB717BD0EFFF8FF4CB5FEFB59FAA1" blockId="11.[151,1437,1050,2013]" pageId="11" pageNumber="162">
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Typical of
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<taxonomicName id="4C12CC94BD0EFFF8FEC9B5FEFE16FAF9" box="[322,435,1267,1293]" class="Clitellata" family="Glossiphoniidae" genus="Helobdella" kingdom="Animalia" order="Rhynchobdellida" pageId="11" pageNumber="162" phylum="Annelida" rank="species" species="lineata">
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<emphasis id="B9666B05BD0EFFF8FEC9B5FEFE16FAF9" box="[322,435,1267,1293]" italics="true" pageId="11" pageNumber="162">H. lineata</emphasis>
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</taxonomicName>
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, all twelve individuals in the archetype series had quickly faded in the preservative with the consequence that pigmentation is generally lost. The pigment pattern described in this section is based primarily on additional specimens photographed in life (HL-24, HL-26)(
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<figureCitation id="1329AB92BD0EFFF8FCDCB436FC3CFAA1" box="[855,921,1339,1365]" captionStart="FIGURE 6" captionStartId="8.[151,250,1824,1848]" captionTargetBox="[362,1231,1027,1798]" captionTargetId="figure-368@8.[356,1231,1027,1798]" captionTargetPageId="8" captionText="FIGURE 6. Variation in dorsal pigmentation and papillation in H. lineata. Two individuals collected from the same location. A, (HL-4), lightly pigmented with metameric spots. B, (HL-26), darkly pigmented with metameric spots. By far the majority of individuals throughout the Albemarle region were lightly pigmented with metameric white dots as illustrated in A. Note: pigmentation disappeared quickly after preservation making the specimen translucent. Abbreviations: b, brooding depression on ventral side; e, eye; m, medial stripe; p, posterior crop caecum; rectum; pa, papilla; ws, metameric white spot. Scale bar: 1.0 mm" figureDoi="http://doi.org/10.5281/zenodo.11233704" httpUri="https://zenodo.org/record/11233704/files/figure.png" pageId="11" pageNumber="162">Fig. 6</figureCitation>
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) or recorded at time of capture.
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</paragraph>
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<paragraph id="8BADB717BD0EFFF8FF1CB452FECFF9FD" blockId="11.[151,1437,1050,2013]" pageId="11" pageNumber="162">
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Size Inexplicably, the three largest individuals of
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<taxonomicName id="4C12CC94BD0EFFF8FD4AB452FC97FA8D" box="[705,818,1375,1401]" class="Clitellata" family="Glossiphoniidae" genus="Helobdella" kingdom="Animalia" order="Rhynchobdellida" pageId="11" pageNumber="162" phylum="Annelida" rank="species" species="lineata">
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<emphasis id="B9666B05BD0EFFF8FD4AB452FC97FA8D" box="[705,818,1375,1401]" italics="true" pageId="11" pageNumber="162">H. lineata</emphasis>
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</taxonomicName>
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(18.5, 16.5 and
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<quantity id="4CEA1AF2BD0EFFF8FC63B453FBEBFA8D" box="[1000,1102,1374,1401]" metricMagnitude="-2" metricUnit="m" metricValue="1.55" pageId="11" pageNumber="162" unit="mm" value="15.5">15.5 mm</quantity>
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, respectively) collected in the 12 years of this study came from precisely the same locality, namely Williams Mill Pond in Wayne County (HL-6). In contrast, the leeches collected in the rest of the Albemarle region were significantly smaller (mean,
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||
<quantity id="4CEA1AF2BD0EFFF8FAB5B4AAFA3DFA35" box="[1342,1432,1447,1473]" metricMagnitude="-3" metricUnit="m" metricValue="7.2" pageId="11" pageNumber="162" unit="mm" value="7.2">7.2 mm</quantity>
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; range,
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<quantity id="4CEA1AF2BD0EFFF8FF68B4C7FEDFFA11" box="[227,378,1482,1509]" metricMagnitude="-3" metricUnit="m" metricValue="7.35" metricValueMax="12.5" metricValueMin="2.2" pageId="11" pageNumber="162" unit="mm" value="7.35" valueMax="12.5" valueMin="2.2">2.2–12.5 mm</quantity>
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; N = 63), with exception of one large individual (
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<quantity id="4CEA1AF2BD0EFFF8FC2BB4C7FBA0FA11" box="[928,1029,1482,1509]" metricMagnitude="-2" metricUnit="m" metricValue="1.55" pageId="11" pageNumber="162" unit="mm" value="15.5">15.5 mm</quantity>
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) collected in a diked canal in Tyrrell County (HL-15A).
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</paragraph>
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<paragraph id="8BADB717BD0EFFF8FF4CB71EFBAEF981" blockId="11.[151,1437,1050,2013]" pageId="11" pageNumber="162">
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Body shape Living individuals of
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<taxonomicName id="4C12CC94BD0EFFF8FDB4B71EFD0BF9D9" box="[575,686,1555,1581]" class="Clitellata" family="Glossiphoniidae" genus="Helobdella" kingdom="Animalia" order="Rhynchobdellida" pageId="11" pageNumber="162" phylum="Annelida" rank="species" species="lineata">
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<emphasis id="B9666B05BD0EFFF8FDB4B71EFD0BF9D9" box="[575,686,1555,1581]" italics="true" pageId="11" pageNumber="162">H. lineata</emphasis>
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</taxonomicName>
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are generally flat.At rest they are widest in the posterior third of the body, and the head and oral sucker are somewhat rounded, not pointed. Interestingly, preserved individuals change body shape, becoming noticeably more narrow and rounded after preservation.
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</paragraph>
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<paragraph id="8BADB717BD0EFFF8FF4CB772FBF5F8B9" blockId="11.[151,1437,1050,2013]" pageId="11" pageNumber="162">
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The reference specimen (HL-5A) is straight (
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||
<quantity id="4CEA1AF2BD0EFFF8FD42B772FC94F96D" box="[713,817,1663,1689]" metricMagnitude="-2" metricUnit="m" metricValue="1.28" pageId="11" pageNumber="162" unit="mm" value="12.8">12.8 mm</quantity>
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) and well preserved but its pigment has faded entirely (
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||
<figureCitation id="1329AB92BD0EFFF8FF14B7AEFF43F949" box="[159,230,1699,1725]" captionStart="FIGURE 4" captionStartId="5.[151,250,1904,1928]" captionTargetBox="[299,1288,774,1878]" captionTargetId="figure-275@5.[299,1288,774,1878]" captionTargetPageId="5" captionText="FIGURE 4. The archetype or reference specimen (HL-5A) on which the description of H. lineata is based. This specimen has faded and become translucent following preservation with 5% formalin. The type locality is in Pasquotank County, NC [36.191176, -76.228516] A, dorsal view. B, ventral view. Abbreviations: a, atrium; cs, caudal sucker; ic, intestinal caecum; pc, post caecum; pr, proboscis; r, rectum; rb, rectal bladder; sv, seminal vesicle; tp, tip of proboscis. Scale bar: 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.11233700" httpUri="https://zenodo.org/record/11233700/files/figure.png" pageId="11" pageNumber="162">Fig. 4</figureCitation>
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). Consequentially, the proboscis (length,
|
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<quantity id="4CEA1AF2BD0EFFF8FD30B7AEFCB6F949" box="[699,787,1699,1725]" metricMagnitude="-3" metricUnit="m" metricValue="2.8" pageId="11" pageNumber="162" unit="mm" value="2.8">2.8 mm</quantity>
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) and atrium are prominent landmarks visible through the translucent body wall. The maximum body width (
|
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<quantity id="4CEA1AF2BD0EFFF8FD63B7CAFCE1F915" box="[744,836,1735,1761]" metricMagnitude="-3" metricUnit="m" metricValue="3.3" pageId="11" pageNumber="162" unit="mm" value="3.3">3.3 mm</quantity>
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) is located toward the posterior end of the body at about segment XIX. From this point the body narrows anteriorly slightly to about segment XI. From here the body narrows increasingly to the oral sucker (width,
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<quantity id="4CEA1AF2BD0EFFF8FD2EB602FD5AF8DD" box="[677,767,1807,1833]" metricMagnitude="-4" metricUnit="m" metricValue="8.0" pageId="11" pageNumber="162" unit="mm" value="0.8">0.8 mm</quantity>
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||
). The distance from the atrium (male gonopore) to the tip of the oral sucker is
|
||
<quantity id="4CEA1AF2BD0EFFF8FED6B63EFE10F8B9" box="[349,437,1843,1869]" metricMagnitude="-3" metricUnit="m" metricValue="3.8" pageId="11" pageNumber="162" unit="mm" value="3.8">3.8 mm</quantity>
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||
, and the width of the body at the male gonopore is
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||
<quantity id="4CEA1AF2BD0EFFF8FC78B63EFBE9F8B9" box="[1011,1100,1843,1869]" metricMagnitude="-3" metricUnit="m" metricValue="2.1" pageId="11" pageNumber="162" unit="mm" value="2.1">2.1 mm</quantity>
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||
.
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||
</paragraph>
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<paragraph id="8BADB717BD0EFFF8FF4CB65AFBA5F861" blockId="11.[151,1437,1050,2013]" pageId="11" pageNumber="162">Gravid and brooding individuals are recognisable because the lateral margins of the body curve downward and inward to form a slight protective cavity. This swells the line of the mid-body.</paragraph>
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<paragraph id="8BADB717BD0EFFF8FF4CB692FA8EF829" blockId="11.[151,1437,1050,2013]" pageId="11" pageNumber="162">
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Oral sucker and mouth Dorsally the oral sucker is almost indistinguishable from the rounded head. From a ventral perspective, however, the oral sucker is clearly outlined as a broad triangular rim (0.5 x
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<quantity id="4CEA1AF2BD0EFFF8FB4EB6CEFABAF829" box="[1221,1311,1987,2013]" metricMagnitude="-4" metricUnit="m" metricValue="9.0" pageId="11" pageNumber="162" unit="mm" value="0.9">0.9 mm</quantity>
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||
).
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||
</paragraph>
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<paragraph id="8BADB717BD09FFFFFF4CB19AFA2DFE7D" blockId="12.[151,1437,151,717]" pageId="12" pageNumber="163">
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The mouth of
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<taxonomicName id="4C12CC94BD09FFFFFEE2B19AFE7FFF45" box="[361,474,151,177]" class="Clitellata" family="Glossiphoniidae" genus="Helobdella" kingdom="Animalia" order="Rhynchobdellida" pageId="12" pageNumber="163" phylum="Annelida" rank="species" species="lineata">
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<emphasis id="B9666B05BD09FFFFFEE2B19AFE7FFF45" box="[361,474,151,177]" italics="true" pageId="12" pageNumber="163">H. lineata</emphasis>
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</taxonomicName>
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differs from most other glossiphoniids in being a conspicuous slit located noticeably closer to the centre of the oral sucker than to the sucker apex. This peculiar anatomy probably reflects the unusual feeding habits of this snail-feeding leech. In this context, it is proposed that the open slit functions as a distensible jaw, capable of receiving the relatively large proboscis when the latter is everted (
|
||
<figureCitation id="1329AB92BD09FFFFFBB1B00EFB01FEE9" box="[1082,1188,259,285]" captionStart="FIGURE 12" captionStartId="15.[151,250,465,489]" captionTargetBox="[151,1436,179,439]" captionTargetId="figure-16@15.[151,1436,179,439]" captionTargetPageId="15" captionText="FIGURE 12. Terminal structure of the proboscis of H. lineata at increasing magnifications. A (HL-9L), ventral view. Everted proboscis showing unusually large mouth; tip of proboscis with circular array of lobules; the proboscis and its lumen widens near the tip. B, C, salivary ductules and close-up of lobules at the proboscis tip; D, salivary pores with secretions at tip of proboscis. Abbreviations: lu, lumen of proboscis; lt, lobule of tip of proboscis; mo, mouth; os, oral sucker; ot, opening of tip of proboscis; sd, salivary ductule; sp, salivary pores at tip of proboscis. Scale bars: A, 1.0 mm; C, 0.1 mm." figureDoi="http://doi.org/10.5281/zenodo.11233719" httpUri="https://zenodo.org/record/11233719/files/figure.png" pageId="12" pageNumber="163">Fig. 12A</figureCitation>
|
||
). In fact, as discussed elsewhere, the proboscis can expand further upon imbibing fluid from a pre-digested snail. Structure of the mouthproboscis complex of this species is of considerable taxonomic significance and contrasts with the small pore located very near the apex of the oral sucker as found in the sanguivorous allied species,
|
||
<taxonomicName id="4C12CC94BD09FFFFFBF0B062FB59FE7D" authorityName="Filippi" authorityYear="1849" box="[1147,1276,367,393]" class="Clitellata" family="Glossiphoniidae" genus="Haementeria" kingdom="Animalia" order="Rhynchobdellida" pageId="12" pageNumber="163" phylum="Annelida" rank="species" species="ghilianii">
|
||
<emphasis id="B9666B05BD09FFFFFBF0B062FB59FE7D" box="[1147,1276,367,393]" italics="true" pageId="12" pageNumber="163">H. ghilianii</emphasis>
|
||
</taxonomicName>
|
||
(pers. obs.).
|
||
</paragraph>
|
||
<paragraph id="8BADB717BD09FFFFFF4CB09EFD73FDED" blockId="12.[151,1437,151,717]" pageId="12" pageNumber="163">Mouth-like slits occur in virtually all specimens of this species collected in the Albemarle region and is considered a non-variable character for taxonomic purposes. Most individuals have closed slits, but the mouth of the reference specimen (HL-5A) is slightly open to reveal a puckered inner lining. Open slits are found in a few other individuals (for example, HL-13A, HL-15A).</paragraph>
|
||
<paragraph id="8BADB717BD09FFFFFF4CB32EFF45FD39" blockId="12.[151,1437,151,717]" pageId="12" pageNumber="163">
|
||
Eyes As a rule
|
||
<taxonomicName id="4C12CC94BD09FFFFFEFEB32EFE43FDC9" box="[373,486,547,573]" class="Clitellata" family="Glossiphoniidae" genus="Helobdella" kingdom="Animalia" order="Rhynchobdellida" pageId="12" pageNumber="163" phylum="Annelida" rank="species" species="lineata">
|
||
<emphasis id="B9666B05BD09FFFFFEFEB32EFE43FDC9" box="[373,486,547,573]" italics="true" pageId="12" pageNumber="163">H. lineata</emphasis>
|
||
</taxonomicName>
|
||
has two and only two well-separated eyes (
|
||
<figureCitation id="1329AB92BD09FFFFFC52B32EFBEAFDC9" box="[985,1103,547,573]" captionStart="FIGURE 9" captionStartId="12.[151,250,1302,1326]" captionTargetBox="[158,1429,738,1275]" captionTargetId="figure-282@12.[158,1429,738,1275]" captionTargetPageId="12" captionText="FIGURE 9. Taxonomic significance of eyes of Helobdella compared to Placobdella. A, B, Helobdella lineata; C, Placobdella multilineata. A, Distance between eyes is maximal when the proboscis (pr) is everted. B, When the proboscis is retained internally the eyes are closer together. C, In the turtle leech P. multilineata the eyes coalesce physically at the mid-line and are not moveable relative to each other. Furthermore, each apparent ‘eye’ of P. multilineata on each side is composite, being comprised of a larger anterior eye and a smaller posterior eye. Scale bars: A, B, 0.3 mm; C, 0.1 mm." figureDoi="http://doi.org/10.5281/zenodo.11233712" httpUri="https://zenodo.org/record/11233712/files/figure.png" pageId="12" pageNumber="163">Fig. 9A,B</figureCitation>
|
||
). This is also the case for the sympatric congener species
|
||
<taxonomicName id="4C12CC94BD09FFFFFE59B34AFDFCFD95" baseAuthorityName="Linnaeus" baseAuthorityYear="1758" box="[466,601,583,609]" class="Clitellata" family="Glossiphoniidae" genus="Helobdella" kingdom="Animalia" order="Rhynchobdellida" pageId="12" pageNumber="163" phylum="Annelida" rank="species" species="stagnalis">
|
||
<emphasis id="B9666B05BD09FFFFFE59B34AFDFCFD95" box="[466,601,583,609]" italics="true" pageId="12" pageNumber="163">H. stagnalis</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C12CC94BD09FFFFFD04B34AFCB1FD95" baseAuthorityName="Castle" baseAuthorityYear="1900" box="[655,788,583,609]" class="Clitellata" family="Glossiphoniidae" genus="Helobdella" kingdom="Animalia" order="Rhynchobdellida" pageId="12" pageNumber="163" phylum="Annelida" rank="species" species="elongata">
|
||
<emphasis id="B9666B05BD09FFFFFD04B34AFCB1FD95" box="[655,788,583,609]" italics="true" pageId="12" pageNumber="163">H. elongata</emphasis>
|
||
</taxonomicName>
|
||
, and is considered characteristic of the genera
|
||
<taxonomicName id="4C12CC94BD09FFFFFAABB34AFE22FD71" authority="(Blanchard, R. 1896)" baseAuthorityName="Blanchard, R." baseAuthorityYear="1896" class="Clitellata" family="Glossiphoniidae" genus="Helobdella" kingdom="Animalia" order="Rhynchobdellida" pageId="12" pageNumber="163" phylum="Annelida" rank="genus">
|
||
<emphasis id="B9666B05BD09FFFFFAABB34AFA39FD95" box="[1312,1436,583,609]" italics="true" pageId="12" pageNumber="163">Helobdella</emphasis>
|
||
(Blanchard, R. 1896)
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C12CC94BD09FFFFFE4AB366FCA7FD71" authority="(Filippi, 1849)" baseAuthorityName="Filippi" baseAuthorityYear="1849" box="[449,770,619,645]" class="Clitellata" family="Glossiphoniidae" genus="Haementeria" kingdom="Animalia" order="Rhynchobdellida" pageId="12" pageNumber="163" phylum="Annelida" rank="genus">
|
||
<emphasis id="B9666B05BD09FFFFFE4AB366FDF6FD70" box="[449,595,619,644]" italics="true" pageId="12" pageNumber="163">Haementeria</emphasis>
|
||
(
|
||
<bibRefCitation id="EF83CAE6BD09FFFFFDEFB366FD5CFD71" author="Filippi, F. de" box="[612,761,619,645]" pageId="12" pageNumber="163" pagination="1 - 14" refId="ref17716" refString="Filippi, F. de (1849) Sopra un nuovo genere (Haementeria) di annelidi della famiglia delle sanguisughe. Memorie della Reale Accademia delle Scienze di Torino, 2 (10), 1 - 14." type="journal article" year="1849">Filippi, 1849</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
(
|
||
<bibRefCitation id="EF83CAE6BD09FFFFFC99B366FC17FD71" author="Sawyer, R." box="[786,946,619,645]" pageId="12" pageNumber="163" refId="ref19495" refString="Sawyer, R. T (1986) Leech Biology and Behaviour. Oxford University Press, Oxford, 1065 pp." type="book" year="1986">Sawyer, 1986</bibRefCitation>
|
||
, 654). In this context the reader is referred to an informative study on the underlying genetics, development and evolution of
|
||
<taxonomicName id="4C12CC94BD09FFFFFBCDB382FB67FD5D" box="[1094,1218,655,681]" class="Clitellata" family="Glossiphoniidae" genus="Helobdella" kingdom="Animalia" order="Rhynchobdellida" pageId="12" pageNumber="163" phylum="Annelida" rank="genus">
|
||
<emphasis id="B9666B05BD09FFFFFBCDB382FB67FD5D" box="[1094,1218,655,681]" italics="true" pageId="12" pageNumber="163">Helobdella</emphasis>
|
||
</taxonomicName>
|
||
eyes (Kwak,
|
||
<emphasis id="B9666B05BD09FFFFFAE9B382FA38FD5D" box="[1378,1437,655,681]" italics="true" pageId="12" pageNumber="163">et al.</emphasis>
|
||
2023).
|
||
</paragraph>
|
||
<caption id="DF6DE79FBD09FFFFFF1CB41BFB2BFA49" ID-DOI="http://doi.org/10.5281/zenodo.11233712" ID-Zenodo-Dep="11233712" httpUri="https://zenodo.org/record/11233712/files/figure.png" pageId="12" pageNumber="163" startId="12.[151,250,1302,1326]" targetBox="[158,1429,738,1275]" targetPageId="12" targetType="figure">
|
||
<paragraph id="8BADB717BD09FFFFFF1CB41BFB2BFA49" blockId="12.[151,1437,1302,1470]" pageId="12" pageNumber="163">
|
||
<emphasis id="B9666B05BD09FFFFFF1CB41BFEB7FADA" bold="true" box="[151,274,1302,1326]" pageId="12" pageNumber="163">FIGURE 9.</emphasis>
|
||
Taxonomic significance of eyes of
|
||
<taxonomicName id="4C12CC94BD09FFFFFDF2B41BFD4FFADA" box="[633,746,1302,1326]" class="Clitellata" family="Glossiphoniidae" genus="Helobdella" kingdom="Animalia" order="Rhynchobdellida" pageId="12" pageNumber="163" phylum="Annelida" rank="genus">
|
||
<emphasis id="B9666B05BD09FFFFFDF2B41BFD4FFADA" box="[633,746,1302,1326]" italics="true" pageId="12" pageNumber="163">Helobdella</emphasis>
|
||
</taxonomicName>
|
||
compared to
|
||
<taxonomicName id="4C12CC94BD09FFFFFCF8B41BFC48FADA" authorityName="R.Blanchard" authorityYear="1893" box="[883,1005,1302,1326]" class="Clitellata" family="Glossiphoniidae" genus="Placobdella" kingdom="Animalia" order="Rhynchobdellida" pageId="12" pageNumber="163" phylum="Annelida" rank="genus">
|
||
<emphasis id="B9666B05BD09FFFFFCF8B41BFC48FADA" box="[883,1005,1302,1326]" italics="true" pageId="12" pageNumber="163">Placobdella</emphasis>
|
||
</taxonomicName>
|
||
. A, B,
|
||
<taxonomicName id="4C12CC94BD09FFFFFBBEB41BFB54FADA" box="[1077,1265,1302,1326]" class="Clitellata" family="Glossiphoniidae" genus="Helobdella" kingdom="Animalia" order="Rhynchobdellida" pageId="12" pageNumber="163" phylum="Annelida" rank="species" species="lineata">
|
||
<emphasis id="B9666B05BD09FFFFFBBEB41BFB54FADA" box="[1077,1265,1302,1326]" italics="true" pageId="12" pageNumber="163">Helobdella lineata</emphasis>
|
||
</taxonomicName>
|
||
; C,
|
||
<taxonomicName id="4C12CC94BD09FFFFFAA9B41BFEB4FAA6" authorityName="Moore" authorityYear="1953" class="Clitellata" family="Glossiphoniidae" genus="Placobdella" kingdom="Animalia" order="Rhynchobdellida" pageId="12" pageNumber="163" phylum="Annelida" rank="species" species="multilineata">
|
||
<emphasis id="B9666B05BD09FFFFFAA9B41BFEB4FAA6" italics="true" pageId="12" pageNumber="163">Placobdella multilineata</emphasis>
|
||
</taxonomicName>
|
||
. A, Distance between eyes is maximal when the proboscis (pr) is everted. B, When the proboscis is retained internally the eyes are closer together. C, In the turtle leech
|
||
<taxonomicName id="4C12CC94BD09FFFFFC87B452FC04FA82" authorityName="Moore" authorityYear="1953" box="[780,929,1374,1398]" class="Clitellata" family="Glossiphoniidae" genus="Placobdella" kingdom="Animalia" order="Rhynchobdellida" pageId="12" pageNumber="163" phylum="Annelida" rank="species" species="multilineata">
|
||
<emphasis id="B9666B05BD09FFFFFC87B452FC04FA82" box="[780,929,1374,1398]" italics="true" pageId="12" pageNumber="163">P. multilineata</emphasis>
|
||
</taxonomicName>
|
||
the eyes coalesce physically at the mid-line and are not moveable relative to each other. Furthermore, each apparent ‘eye’ of
|
||
<taxonomicName id="4C12CC94BD09FFFFFC3BB48EFBE1FA6E" authorityName="Moore" authorityYear="1953" box="[944,1092,1410,1434]" class="Clitellata" family="Glossiphoniidae" genus="Placobdella" kingdom="Animalia" order="Rhynchobdellida" pageId="12" pageNumber="163" phylum="Annelida" rank="species" species="multilineata">
|
||
<emphasis id="B9666B05BD09FFFFFC3BB48EFBE1FA6E" box="[944,1092,1410,1434]" italics="true" pageId="12" pageNumber="163">P. multilineata</emphasis>
|
||
</taxonomicName>
|
||
on each side is composite, being comprised of a larger anterior eye and a smaller posterior eye. Scale bars: A, B, 0.3 mm; C, 0.1 mm.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="8BADB717BD09FFFFFF4CB71DFB66F8BE" blockId="12.[151,1437,1552,2011]" pageId="12" pageNumber="163">
|
||
The reference specimen has no visible eyes, as is the case for many of the faded specimens in this collection but in a comparable specimen (HL-15A) a single pair of well-separated eyes lies in the posterior part of segment III (i.e. on annulus III (a2+a3) (
|
||
<figureCitation id="1329AB92BD09FFFFFE17B755FDA4F986" box="[412,513,1624,1650]" captionStart="FIGURE 10" captionStartId="13.[151,250,991,1015]" captionTargetBox="[153,1432,342,964]" captionTargetId="figure-92@13.[153,1435,342,965]" captionTargetPageId="13" captionText="FIGURE 10. Evidence that the neck region of H. lineata is capable of extending and swelling (see section on feeding biology). A, proboscis everted; B–D, proboscis at rest, i.e. not everted. A (HL-9D), neck region narrows (white arrow) when proboscis is extended; B (HL-15A), annulation of the neck region at rest is more subdivided than annuli in mid-body region; C (HL-13A), ventral nerve cord (blue) between ganglia VIII–IX and IX–X is characteristically sinuous (black arrow) when individual is at rest. D (HL-13A), single salivary ductules are sinuous (black arrow) within the proboscis when individual is at rest. Abbreviations: co, circumpharyngeal commissure (brain); sd, single salivary ductule (brown); sb, salivary bundle (brown). Scale bar: 10 mm." figureDoi="http://doi.org/10.5281/zenodo.11233714" httpUri="https://zenodo.org/record/11233714/files/figure.png" pageId="12" pageNumber="163">Fig. 10B</figureCitation>
|
||
). Functionally, the distance between the eyes is variable in
|
||
<taxonomicName id="4C12CC94BD09FFFFFB00B754FB5FF986" box="[1163,1274,1624,1650]" class="Clitellata" family="Glossiphoniidae" genus="Helobdella" kingdom="Animalia" order="Rhynchobdellida" pageId="12" pageNumber="163" phylum="Annelida" rank="species" species="lineata">
|
||
<emphasis id="B9666B05BD09FFFFFB00B754FB5FF986" box="[1163,1274,1624,1650]" italics="true" pageId="12" pageNumber="163">H. lineata</emphasis>
|
||
</taxonomicName>
|
||
but they never adjoin at the midline. A previously unreported observation in
|
||
<taxonomicName id="4C12CC94BD09FFFFFCCCB771FC66F962" box="[839,963,1660,1686]" class="Clitellata" family="Glossiphoniidae" genus="Helobdella" kingdom="Animalia" order="Rhynchobdellida" pageId="12" pageNumber="163" phylum="Annelida" rank="genus">
|
||
<emphasis id="B9666B05BD09FFFFFCCCB771FC66F962" box="[839,963,1660,1686]" italics="true" pageId="12" pageNumber="163">Helobdella</emphasis>
|
||
</taxonomicName>
|
||
is that when the relatively large proboscis is everted the distance between the eyes adjusts spatially. In other words distance between eyes is maximal when the proboscis is everted (
|
||
<figureCitation id="1329AB92BD09FFFFFE27B7C9FDA3F92A" box="[428,518,1732,1758]" captionStart="FIGURE 9" captionStartId="12.[151,250,1302,1326]" captionTargetBox="[158,1429,738,1275]" captionTargetId="figure-282@12.[158,1429,738,1275]" captionTargetPageId="12" captionText="FIGURE 9. Taxonomic significance of eyes of Helobdella compared to Placobdella. A, B, Helobdella lineata; C, Placobdella multilineata. A, Distance between eyes is maximal when the proboscis (pr) is everted. B, When the proboscis is retained internally the eyes are closer together. C, In the turtle leech P. multilineata the eyes coalesce physically at the mid-line and are not moveable relative to each other. Furthermore, each apparent ‘eye’ of P. multilineata on each side is composite, being comprised of a larger anterior eye and a smaller posterior eye. Scale bars: A, B, 0.3 mm; C, 0.1 mm." figureDoi="http://doi.org/10.5281/zenodo.11233712" httpUri="https://zenodo.org/record/11233712/files/figure.png" pageId="12" pageNumber="163">Fig. 9A</figureCitation>
|
||
), unlike when the proboscis is at rest, internally (
|
||
<figureCitation id="1329AB92BD09FFFFFBA3B7C9FB25F92A" box="[1064,1152,1732,1758]" captionStart="FIGURE 9" captionStartId="12.[151,250,1302,1326]" captionTargetBox="[158,1429,738,1275]" captionTargetId="figure-282@12.[158,1429,738,1275]" captionTargetPageId="12" captionText="FIGURE 9. Taxonomic significance of eyes of Helobdella compared to Placobdella. A, B, Helobdella lineata; C, Placobdella multilineata. A, Distance between eyes is maximal when the proboscis (pr) is everted. B, When the proboscis is retained internally the eyes are closer together. C, In the turtle leech P. multilineata the eyes coalesce physically at the mid-line and are not moveable relative to each other. Furthermore, each apparent ‘eye’ of P. multilineata on each side is composite, being comprised of a larger anterior eye and a smaller posterior eye. Scale bars: A, B, 0.3 mm; C, 0.1 mm." figureDoi="http://doi.org/10.5281/zenodo.11233712" httpUri="https://zenodo.org/record/11233712/files/figure.png" pageId="12" pageNumber="163">Fig. 9B</figureCitation>
|
||
). In contrast the eyes of a sympatric turtle leech
|
||
<taxonomicName id="4C12CC94BD09FFFFFE04B7E4FD94F8F6" authorityName="Moore" authorityYear="1953" box="[399,561,1768,1794]" class="Clitellata" family="Glossiphoniidae" genus="Placobdella" kingdom="Animalia" order="Rhynchobdellida" pageId="12" pageNumber="163" phylum="Annelida" rank="species" species="multilineata">
|
||
<emphasis id="B9666B05BD09FFFFFE04B7E4FD94F8F6" box="[399,561,1768,1794]" italics="true" pageId="12" pageNumber="163">P. multilineata</emphasis>
|
||
</taxonomicName>
|
||
coalesce physically at the mid-line and are not moveable relative to each other (
|
||
<figureCitation id="1329AB92BD09FFFFFF14B601FF52F8D2" box="[159,247,1804,1830]" captionStart="FIGURE 9" captionStartId="12.[151,250,1302,1326]" captionTargetBox="[158,1429,738,1275]" captionTargetId="figure-282@12.[158,1429,738,1275]" captionTargetPageId="12" captionText="FIGURE 9. Taxonomic significance of eyes of Helobdella compared to Placobdella. A, B, Helobdella lineata; C, Placobdella multilineata. A, Distance between eyes is maximal when the proboscis (pr) is everted. B, When the proboscis is retained internally the eyes are closer together. C, In the turtle leech P. multilineata the eyes coalesce physically at the mid-line and are not moveable relative to each other. Furthermore, each apparent ‘eye’ of P. multilineata on each side is composite, being comprised of a larger anterior eye and a smaller posterior eye. Scale bars: A, B, 0.3 mm; C, 0.1 mm." figureDoi="http://doi.org/10.5281/zenodo.11233712" httpUri="https://zenodo.org/record/11233712/files/figure.png" pageId="12" pageNumber="163">Fig. 9C</figureCitation>
|
||
). The immutability of the distance between eyes in the turtle leech may be attributable in part to the lack of need for spatial adjustment required by its long slender proboscis in
|
||
<taxonomicName id="4C12CC94BD09FFFFFC1EB63CFB9DF8BE" authorityName="Moore" authorityYear="1953" box="[917,1080,1840,1866]" class="Clitellata" family="Glossiphoniidae" genus="Placobdella" kingdom="Animalia" order="Rhynchobdellida" pageId="12" pageNumber="163" phylum="Annelida" rank="species" species="multilineata">
|
||
<emphasis id="B9666B05BD09FFFFFC1EB63CFB9DF8BE" box="[917,1080,1840,1866]" italics="true" pageId="12" pageNumber="163">P. multilineata</emphasis>
|
||
</taxonomicName>
|
||
(pers. obs.).
|
||
</paragraph>
|
||
<paragraph id="8BADB717BD09FFFFFF4CB659FAD8F82F" blockId="12.[151,1437,1552,2011]" pageId="12" pageNumber="163">
|
||
Caudal sucker The caudal sucker of the reference specimen (HL-5A) is almost circular (
|
||
<quantity id="4CEA1AF2BD09FFFFFB19B659FB4EF89A" box="[1170,1259,1876,1903]" metricMagnitude="-3" metricUnit="m" metricValue="1.5" pageId="12" pageNumber="163" unit="mm" value="1.5">1.5 mm</quantity>
|
||
). In lateral view the posterior end of the body has a pronounced concave curve to meet the caudal sucker. This curve rides high over the centre of the sucker such that upon dorsal view only a small portion (
|
||
<quantity id="4CEA1AF2BD09FFFFFC4FB691FB8FF842" box="[964,1066,1948,1974]" metricMagnitude="-4" metricUnit="m" metricValue="2.4" pageId="12" pageNumber="163" unit="mm" value="0.24">0.24 mm</quantity>
|
||
) of the caudal sucker is exposed. This peculiar, somewhat humped, configuration may reflect the state of contents of the underlying large rectum.
|
||
</paragraph>
|
||
<paragraph id="8BADB717BD08FFFEFF4CB19AFE3CFEB5" blockId="13.[151,1437,151,321]" pageId="13" pageNumber="164">
|
||
Annulation In the reference specimen annuli of the neck region are irregularly sub-divided in keeping with extensibility of the neck (
|
||
<figureCitation id="1329AB92BD08FFFEFE39B1B6FD93FF21" box="[434,566,187,213]" captionStart="FIGURE 10" captionStartId="13.[151,250,991,1015]" captionTargetBox="[153,1432,342,964]" captionTargetId="figure-92@13.[153,1435,342,965]" captionTargetPageId="13" captionText="FIGURE 10. Evidence that the neck region of H. lineata is capable of extending and swelling (see section on feeding biology). A, proboscis everted; B–D, proboscis at rest, i.e. not everted. A (HL-9D), neck region narrows (white arrow) when proboscis is extended; B (HL-15A), annulation of the neck region at rest is more subdivided than annuli in mid-body region; C (HL-13A), ventral nerve cord (blue) between ganglia VIII–IX and IX–X is characteristically sinuous (black arrow) when individual is at rest. D (HL-13A), single salivary ductules are sinuous (black arrow) within the proboscis when individual is at rest. Abbreviations: co, circumpharyngeal commissure (brain); sd, single salivary ductule (brown); sb, salivary bundle (brown). Scale bar: 10 mm." figureDoi="http://doi.org/10.5281/zenodo.11233714" httpUri="https://zenodo.org/record/11233714/files/figure.png" pageId="13" pageNumber="164">Figure 10B</figureCitation>
|
||
). On the other hand annuli of the mid-body region are more evenly expressed, but these too are occasionally subdivided. Often in some specimens (HL-5G) the subdivision is in the posterior third of the annulus rather than in the middle. In other words annular subdivision is an incompletely expressed (variable) character in
|
||
<taxonomicName id="4C12CC94BD08FFFEFEA9B02AFE37FEB5" box="[290,402,295,321]" class="Clitellata" family="Glossiphoniidae" genus="Helobdella" kingdom="Animalia" order="Rhynchobdellida" pageId="13" pageNumber="164" phylum="Annelida" rank="species" species="lineata">
|
||
<emphasis id="B9666B05BD08FFFEFEA9B02AFE37FEB5" box="[290,402,295,321]" italics="true" pageId="13" pageNumber="164">H. lineata</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</paragraph>
|
||
<caption id="DF6DE79FBD08FFFEFF1CB2D2FA30FB5F" ID-DOI="http://doi.org/10.5281/zenodo.11233714" ID-Zenodo-Dep="11233714" httpUri="https://zenodo.org/record/11233714/files/figure.png" pageId="13" pageNumber="164" startId="13.[151,250,991,1015]" targetBox="[153,1432,342,964]" targetPageId="13" targetType="figure">
|
||
<paragraph id="8BADB717BD08FFFEFF1CB2D2FA30FB5F" blockId="13.[151,1437,991,1196]" pageId="13" pageNumber="164">
|
||
<emphasis id="B9666B05BD08FFFEFF1CB2D2FEBAFC0C" bold="true" box="[151,287,991,1016]" pageId="13" pageNumber="164">FIGURE 10.</emphasis>
|
||
Evidence that the neck region of
|
||
<taxonomicName id="4C12CC94BD08FFFEFDF9B2EDFD72FC0C" box="[626,727,992,1016]" class="Clitellata" family="Glossiphoniidae" genus="Helobdella" kingdom="Animalia" order="Rhynchobdellida" pageId="13" pageNumber="164" phylum="Annelida" rank="species" species="lineata">
|
||
<emphasis id="B9666B05BD08FFFEFDF9B2EDFD72FC0C" box="[626,727,992,1016]" italics="true" pageId="13" pageNumber="164">H. lineata</emphasis>
|
||
</taxonomicName>
|
||
is capable of extending and swelling (see section on feeding biology). A, proboscis everted; B–D, proboscis at rest, i.e. not everted. A (HL-9D), neck region narrows (white arrow) when proboscis is extended; B (HL-15A), annulation of the neck region at rest is more subdivided than annuli in mid-body region; C (HL-13A), ventral nerve cord (blue) between ganglia VIII–IX and IX–X is characteristically sinuous (black arrow) when individual is at rest. D (HL-13A), single salivary ductules are sinuous (black arrow) within the proboscis when individual is at rest. Abbreviations: co, circumpharyngeal commissure (brain); sd, single salivary ductule (brown); sb, salivary bundle (brown). Scale bar: 10 mm.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="8BADB717BD08FFFEFF4CB5F3FC08FA5C" blockId="13.[151,1437,1278,1448]" pageId="13" pageNumber="164">
|
||
Incompletely expressed subdivision of annuli also occurs in the two sympatric congener species,
|
||
<taxonomicName id="4C12CC94BD08FFFEFB61B5F2FACAFAEC" baseAuthorityName="Linnaeus" baseAuthorityYear="1758" box="[1258,1391,1278,1304]" class="Clitellata" family="Glossiphoniidae" genus="Helobdella" kingdom="Animalia" order="Rhynchobdellida" pageId="13" pageNumber="164" phylum="Annelida" rank="species" species="stagnalis">
|
||
<emphasis id="B9666B05BD08FFFEFB61B5F2FACAFAEC" box="[1258,1391,1278,1304]" italics="true" pageId="13" pageNumber="164">H. stagnalis</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C12CC94BD08FFFEFF1CB42EFEB9FAC8" baseAuthorityName="Castle" baseAuthorityYear="1900" box="[151,284,1314,1340]" class="Clitellata" family="Glossiphoniidae" genus="Helobdella" kingdom="Animalia" order="Rhynchobdellida" pageId="13" pageNumber="164" phylum="Annelida" rank="species" species="elongata">
|
||
<emphasis id="B9666B05BD08FFFEFF1CB42EFEB9FAC8" box="[151,284,1314,1340]" italics="true" pageId="13" pageNumber="164">H. elongata</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<figureCitation id="1329AB92BD08FFFEFEA0B42FFE3BFAC8" box="[299,414,1314,1340]" captionStart="FIGURE 3" captionStartId="4.[151,250,1864,1888]" captionTargetBox="[322,1265,919,1838]" captionTargetId="figure-328@4.[322,1265,919,1839]" captionTargetPageId="4" captionText="FIGURE 3. Three co-existing species of Helobdella indigenous to the Albemarle region of the Outer Banks, North Carolina. A, H. lineata, the subject of this paper, photographed in life, pigment and papillar pattern intact (HL-26); B, H. stagnalis (sensu lato) (Hst-1A); C, H. elongata (HE-2A). Scale bar: 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.11233698" httpUri="https://zenodo.org/record/11233698/files/figure.png" pageId="13" pageNumber="164">Fig. 3B,C</figureCitation>
|
||
). As a variable character in all three clades annular subdivision is taxonomically unreliable and should not be used to define these or closely allied forms of
|
||
<taxonomicName id="4C12CC94BD08FFFEFC0CB44BFBA6FA94" box="[903,1027,1350,1376]" class="Clitellata" family="Glossiphoniidae" genus="Helobdella" kingdom="Animalia" order="Rhynchobdellida" pageId="13" pageNumber="164" phylum="Annelida" rank="genus">
|
||
<emphasis id="B9666B05BD08FFFEFC0CB44BFBA6FA94" box="[903,1027,1350,1376]" italics="true" pageId="13" pageNumber="164">Helobdella</emphasis>
|
||
</taxonomicName>
|
||
. In this context several ‘species’ of
|
||
<taxonomicName id="4C12CC94BD08FFFEFF1CB467FEB6FA70" box="[151,275,1386,1412]" class="Clitellata" family="Glossiphoniidae" genus="Helobdella" kingdom="Animalia" order="Rhynchobdellida" pageId="13" pageNumber="164" phylum="Annelida" rank="genus">
|
||
<emphasis id="B9666B05BD08FFFEFF1CB467FEB6FA70" box="[151,275,1386,1412]" italics="true" pageId="13" pageNumber="164">Helobdella</emphasis>
|
||
</taxonomicName>
|
||
and even allied ‘genera’ have been defined in terms of subdivided annuli, for example,
|
||
<taxonomicName id="4C12CC94BD08FFFEFB65B466FAD6FA70" box="[1262,1395,1386,1412]" class="Clitellata" family="Glossiphoniidae" genus="Helobdella" kingdom="Animalia" order="Rhynchobdellida" pageId="13" pageNumber="164" phylum="Annelida" rank="species" species="scutifera">
|
||
<emphasis id="B9666B05BD08FFFEFB65B466FAD6FA70" box="[1262,1395,1386,1412]" italics="true" pageId="13" pageNumber="164">H. scutifera</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C12CC94BD08FFFEFA0AB466FEA8FA5C" class="Clitellata" family="Glossiphoniidae" genus="Helobdella" kingdom="Animalia" order="Rhynchobdellida" pageId="13" pageNumber="164" phylum="Annelida" rank="species" species="longicollis">
|
||
<emphasis id="B9666B05BD08FFFEFA0AB466FEA8FA5C" italics="true" pageId="13" pageNumber="164">H. longicollis</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C12CC94BD08FFFEFECFB482FE68FA5C" authorityName="Ringuelet" authorityYear="1948" box="[324,461,1422,1448]" class="Clitellata" family="Glossiphoniidae" genus="Helobdella" kingdom="Animalia" order="Rhynchobdellida" pageId="13" pageNumber="164" phylum="Annelida" rank="species" species="diploides">
|
||
<emphasis id="B9666B05BD08FFFEFECFB482FE68FA5C" box="[324,461,1422,1448]" italics="true" pageId="13" pageNumber="164">H. diploides</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<bibRefCitation id="EF83CAE6BD08FFFEFE50B483FD7BFA5C" author="Siddall, M. E. & Borda, E." box="[475,734,1422,1448]" pageId="13" pageNumber="164" pagination="23 - 33" refId="ref19699" refString="Siddall, M. E. & Borda, E. (2003) Phylogeny and revision of the leech genus Helobdella (Glossiphoniidae) based on mitochondrial gene sequences and morphological data and a special consideration of the triserialis complex. Zoologica Scripta, 32 (1), 23 - 33. https: // doi. org / 10.1046 / j. 1463 - 6409.2003.00098. x" type="journal article" year="2003">Siddall & Borda, 2003</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EF83CAE6BD08FFFEFD62B483FC05FA5C" author="Ringuelet, R. A." box="[745,928,1422,1448]" pageId="13" pageNumber="164" pagination="269 - 276" refId="ref19330" refString="Ringuelet, R. A. (1978) Nuevos generos y especies de Glossiphoniidae Sudamericanos basados en caracteres ecto y endosomaticos (Hirudinea, Glossiphoniiformes). Limnobios, 1, 269 - 276." type="journal article" year="1978">Ringuelet, 1978</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |