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<document id="36FEA18B6E860C373AAEE092B8F2ADF2" ID-DOI="10.5281/zenodo.196223" ID-GBIF-Dataset="648631c9-8b76-4f0f-85a4-d808b6fbfe70" ID-ISSN="1175-5326" ID-Zenodo-Dep="196223" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.taxonomicNames_approvedBy="felipe" checkinTime="1460240568931" checkinUser="plazi" docAuthor="Whittington, Ian D." docDate="2010" docId="03C11D43D31DFFE7FF7F43939F9B5655" docLanguage="en" docName="zt02519p030.pdf" docOrigin="Zootaxa 2519" docStyle="DocumentStyle:890A69B780ED73D6DB8551B71C8AC79E.4:Zootaxa.2009-2012.journal_article" docStyleId="890A69B780ED73D6DB8551B71C8AC79E" docStyleName="Zootaxa.2009-2012.journal_article" docStyleVersion="4" docTitle="Benedeniella macrocolpa (Luhe, 1906) Yamaguti 1963" docType="treatment" docVersion="8" lastPageNumber="15" masterDocId="FFF8653BD318FFE9FFE847349A135202" masterDocTitle="Revision of Benedeniella Johnston, 1929 (Monogenea: Capsalidae), its assignment to Entobdellinae Bychowsky, 1957 and comments on subfamilial composition" masterLastPageNumber="30" masterPageNumber="1" pageNumber="6" updateTime="1698240612236" updateUser="plazi">
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<mods:title id="C9E678A7B81CE640F0F922E28075DEC7">Revision of Benedeniella Johnston, 1929 (Monogenea: Capsalidae), its assignment to Entobdellinae Bychowsky, 1957 and comments on subfamilial composition</mods:title>
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<mods:namePart id="577E5014294B91E52E32373674FEC0C2">Whittington, Ian D.</mods:namePart>
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<mods:date id="4BC50F29E2FF6497224514899D9DBB0A">2010</mods:date>
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<mods:identifier id="2EAF8DB06480ED373F16B0AA9C2C8A27" type="DOI">10.5281/zenodo.196223</mods:identifier>
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<treatment id="03C11D43D31DFFE7FF7F43939F9B5655" ID-DOI="http://doi.org/10.5281/zenodo.6207322" ID-GBIF-Taxon="119389647" ID-Zenodo-Dep="6207322" LSID="urn:lsid:plazi:treatment:03C11D43D31DFFE7FF7F43939F9B5655" httpUri="http://treatment.plazi.org/id/03C11D43D31DFFE7FF7F43939F9B5655" lastPageId="14" lastPageNumber="15" pageId="5" pageNumber="6">
<subSubSection id="C372FFDED31DFFECFF7F43939BE556E6" pageId="5" pageNumber="6" type="nomenclature">
<paragraph id="8BD7AC55D31DFFECFF7F4393992B56C3" blockId="5.[151,824,1191,1252]" box="[151,824,1191,1217]" pageId="5" pageNumber="6">
<heading id="D09F1B39D31DFFECFF7F4393992B56C3" bold="true" box="[151,824,1191,1217]" fontSize="11" level="1" pageId="5" pageNumber="6" reason="1">
<taxonomicName id="4C68D7D6D31DFFECFF7F4393992B56C3" ID-CoL="7SKFK" authority="(Luhe, 1906) Yamaguti, 1963" authorityName="(Luhe, 1906) Yamaguti" authorityYear="1963" box="[151,824,1191,1217]" class="Monogenea" family="Capsalidae" genus="Benedeniella" kingdom="Animalia" order="Capsalidea" pageId="5" pageNumber="6" phylum="Platyhelminthes" rank="species" species="macrocolpa">
<emphasis id="B91C7047D31DFFECFF7F4393992B56C3" bold="true" box="[151,824,1191,1217]" pageId="5" pageNumber="6">
<emphasis id="B91C7047D31DFFECFF7F43939BD956C3" bold="true" box="[151,458,1191,1217]" italics="true" pageId="5" pageNumber="6">Benedeniella macrocolpa</emphasis>
(
<bibRefCitation id="EFF9D1A4D31DFFECFE314393987456C3" author="Luhe" box="[473,615,1191,1217]" pageId="5" pageNumber="29" refString="Luhe, M. (1906) Report on the trematode parasites from the marine fishes of Ceylon. Report to the Government of Ceylon on the Pearl Oyster Fisheries of the Gulf of Manaar, Supplementary Report, 5, 97 - 108." type="journal article" year="1906">Lühe, 1906</bibRefCitation>
)
<bibRefCitation id="EFF9D1A4D31DFFECFD9E4393992B56C3" author="Yamaguti" box="[630,824,1191,1217]" pageId="5" pageNumber="30" refString="Yamaguti, S. (1963) Systema Helminthum. Volume IV. Monogenea and Aspidocotylea. InterScience Publishers, New York, USA, 699 pp." type="book" year="1963">Yamaguti, 1963</bibRefCitation>
</emphasis>
</taxonomicName>
</heading>
</paragraph>
<paragraph id="8BD7AC55D31DFFECFF7F43FE9BE556E6" blockId="5.[151,824,1191,1252]" box="[151,502,1226,1252]" pageId="5" pageNumber="6">
(
<figureCitation id="1353B0D0D31DFFECFF7743FE9AE356E6" box="[159,240,1226,1252]" captionStart="FIGURE 1" captionStartId="6.[151,258,1755,1779]" captionTargetBox="[278,1303,216,1729]" captionTargetId="figure@6.[267,1321,197,1745]" captionTargetPageId="6" captionText="FIGURE 1. Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963. Whole adult parasite, composite drawing, in ventral view based on AHC 29906 (specimen labeled no. 1) with some additions by hand. For clarity, extent of intestinal diverticula (entirely coextensive with the vitellarium) other than immediately posterior to pharynx and separate uterus parallel with terminal male genital duct are not shown and extensive convolutions of vas deferens are simplified (but are shown in full in Figure 6). Abbreviations: aao, anterior attachment organ; ah, anterior hamulus; as, accessory sclerite; c, cirrus; cp, common genital pore; cs, cirrus sac; dh, dorsal horn; e, excretory bladder; em, extrinsic muscle; g, germarium; gr, ventral groove; h, haptor; ho, hooklet; i, intestine; mv, marginal valve; n, longitudinal nerve trunk; o, ootype; p, pharynx; ph, posterior hamulus; rp, rounded papilla; t, testis; te, tendon; v, vagina; vi, vitelline follicle; vp, vaginal pore. Scale bar: 1 mm." httpUri="https://zenodo.org/record/196224/files/figure.png" pageId="5" pageNumber="6">Figs. 1</figureCitation>
,
<figureCitation id="1353B0D0D31DFFECFF1743FE9B0356E6" box="[255,272,1226,1252]" captionStart="FIGURE 2" captionStartId="7.[151,255,1811,1835]" captionTargetBox="[307,1294,216,1780]" captionTargetId="figure@7.[289,1304,194,1795]" captionTargetPageId="7" captionText="FIGURE 2. Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963. Haptoral sclerites of adult parasite. A. Accessory sclerite. B. Anterior hamulus. C. Posterior hamulus. D. Hooklet. E. Proximal ends of anterior hamulus (ah) and accessory sclerite (as) showing main tendon path (stippled) plus branch (solid) to haptor tissue (see also Figure 3 A). Scale bars: A – B, 100 µm; C, 50 µm; D, 10 µm; E, 200 µm." httpUri="https://zenodo.org/record/196225/files/figure.png" pageId="5" pageNumber="6">2</figureCitation>
,
<figureCitation id="1353B0D0D31DFFECFEF443FE9B3A56E6" box="[284,297,1226,1252]" captionStart="FIGURE 3" captionStartId="8.[151,254,1620,1644]" captionTargetBox="[248,1337,531,1594]" captionTargetId="figure@8.[247,1339,530,1595]" captionTargetPageId="8" captionText="FIGURE 3. Differential interference contrast photomicrographs of haptor and anterior attachment organs from stained, mounted specimens of Benedeniella species. A. Accessory sclerite (as) and anterior (= proximal) part of anterior hamulus (ah) on one side of haptor of B. macrocolpa (Lühe, 1906) Yamaguti, 1963 showing route of tendon through proximal notch (pn) of accessory sclerite before attaching to proximal end of anterior hamulus with a branch (b) of the tendon apparently attaching to haptor tissue. AHC 29906 (specimen labeled no. 1). B. The left anterior attachment organ of B. macrocolpa showing complete ventral grooves (gr) and some incomplete grooves (ig) anterolaterally. Note the rounded papillae (rp) at the anterior extremity of the parasite. AHC 29906 (specimen labeled no. 1). C. Accessory sclerite (as) and anterior (= proximal) part of anterior hamulus (ah) on one side of haptor of B. posterocolpa (Hargis, 1955) Yamaguti, 1963 showing route of tendon through proximal notch of accessory sclerite before attaching to proximal end of anterior hamulus with a branch (b) of the tendon apparently attaching to haptor tissue. AHC 29911 (specimen labeled no. 13). D. The left anterior attachment organ of B. posterocolpa showing ventral grooves (gr) and some incomplete grooves (ig) anterolaterally. Note the dorsal horn (dh) where the body and anterior attachment organ join. AHC 29911 (specimen labeled no. 13). Scale bars: 200 µm." httpUri="https://zenodo.org/record/196226/files/figure.png" pageId="5" pageNumber="6">3</figureCitation>
A–B, 4, 5A, 6, 7)
</paragraph>
</subSubSection>
<subSubSection id="C372FFDED31DFFECFF7F42209B17576E" pageId="5" pageNumber="6" type="description">
<paragraph id="8BD7AC55D31DFFECFF7F42209B17576E" blockId="5.[151,1436,1300,1388]" pageId="5" pageNumber="6">
<emphasis id="B91C7047D31DFFECFF7F42209B06572E" bold="true" box="[151,277,1300,1324]" pageId="5" pageNumber="6">Synonyms:</emphasis>
<taxonomicName id="4C68D7D6D31DFFECFEC84221993A572E" authority="Luhe, 1906" authorityName="Luhe" authorityYear="1906" box="[288,809,1301,1324]" class="Monogenea" family="Capsalidae" genus="Epibdella" higherTaxonomySource="CoL" kingdom="Animalia" order="Capsalidea" pageId="5" pageNumber="6" phylum="Platyhelminthes" rank="species" species="macrocolpa" subGenus="Benedenia">
<emphasis id="B91C7047D31DFFECFEC842219B9B572E" box="[288,392,1301,1324]" italics="true" pageId="5" pageNumber="6">Epibdella</emphasis>
(
<emphasis id="B91C7047D31DFFECFE7342219818572E" box="[411,523,1301,1324]" italics="true" pageId="5" pageNumber="6">Benedenia</emphasis>
)
<emphasis id="B91C7047D31DFFECFDC84221988D572E" box="[544,670,1301,1324]" italics="true" pageId="5" pageNumber="6">macrocolpa</emphasis>
<bibRefCitation id="EFF9D1A4D31DFFECFD414221993A572E" author="Luhe" box="[681,809,1301,1324]" pageId="5" pageNumber="29" refString="Luhe, M. (1906) Report on the trematode parasites from the marine fishes of Ceylon. Report to the Government of Ceylon on the Pearl Oyster Fisheries of the Gulf of Manaar, Supplementary Report, 5, 97 - 108." type="journal article" year="1906">Lühe, 1906</bibRefCitation>
</taxonomicName>
;
<taxonomicName id="4C68D7D6D31DFFECFCD142219F8B572E" authority="(Luhe, 1906) MacCallum, 1927" authorityName="(Luhe, 1906) MacCallum" authorityYear="1927" box="[825,1432,1301,1324]" class="Monogenea" family="Capsalidae" genus="Epibdella" higherTaxonomySource="CoL" kingdom="Animalia" order="Capsalidea" pageId="5" pageNumber="6" phylum="Platyhelminthes" rank="species" species="macrocolpa">
<emphasis id="B91C7047D31DFFECFCD142219E39572E" box="[825,1066,1301,1324]" italics="true" pageId="5" pageNumber="6">Epibdella macrocolpa</emphasis>
(
<bibRefCitation id="EFF9D1A4D31DFFECFBD442219EAF572E" author="Luhe" box="[1084,1212,1301,1324]" pageId="5" pageNumber="29" refString="Luhe, M. (1906) Report on the trematode parasites from the marine fishes of Ceylon. Report to the Government of Ceylon on the Pearl Oyster Fisheries of the Gulf of Manaar, Supplementary Report, 5, 97 - 108." type="journal article" year="1906">Lühe, 1906</bibRefCitation>
)
<bibRefCitation id="EFF9D1A4D31DFFECFB2742219F8B572E" author="MacCallum" box="[1231,1432,1301,1324]" pageId="5" pageNumber="29" refString="MacCallum, G. A. (1927) A new ectoparasitic trematode Epibdella melleni, sp. nov. Zoopathologica, 1, 291 - 300." type="journal article" year="1927">MacCallum, 1927</bibRefCitation>
</taxonomicName>
;
<taxonomicName id="4C68D7D6D31DFFECFF2F4201998C574E" authority="(Luhe, 1906) Johnston, 1929" authorityName="(Luhe, 1906) Johnston" authorityYear="1929" box="[199,927,1333,1356]" class="Monogenea" family="Capsalidae" genus="Benedenia" kingdom="Animalia" order="Capsalidea" pageId="5" pageNumber="6" phylum="Platyhelminthes" rank="species" species="macrocolpa" subGenus="Benedeniella">
<emphasis id="B91C7047D31DFFECFF2F42019B2B574E" box="[199,312,1333,1356]" italics="true" pageId="5" pageNumber="6">Benedenia</emphasis>
(
<emphasis id="B91C7047D31DFFECFEAF42019BC1574E" box="[327,466,1333,1356]" italics="true" pageId="5" pageNumber="6">Benedeniella</emphasis>
)
<emphasis id="B91C7047D31DFFECFE0A42019873574E" box="[482,608,1333,1356]" italics="true" pageId="5" pageNumber="6">macrocolpa</emphasis>
(
<bibRefCitation id="EFF9D1A4D31DFFECFD87420198FF574E" author="Luhe" box="[623,748,1333,1356]" pageId="5" pageNumber="29" refString="Luhe, M. (1906) Report on the trematode parasites from the marine fishes of Ceylon. Report to the Government of Ceylon on the Pearl Oyster Fisheries of the Gulf of Manaar, Supplementary Report, 5, 97 - 108." type="journal article" year="1906">Lühe, 1906</bibRefCitation>
)
<bibRefCitation id="EFF9D1A4D31DFFECFD124201998C574E" author="Johnston" box="[762,927,1333,1356]" pageId="5" pageNumber="28" refString="Johnston, T. H. (1929) Remarks on the synonymy of certain tristomatid trematode genera. Transactions of the Royal Society of South Australia, 53, 71 - 78." type="journal article" year="1929">Johnston, 1929</bibRefCitation>
</taxonomicName>
;
<taxonomicName id="4C68D7D6D31DFFECFC4242019B13576E" authority="(Luhe, 1906) Meserve, 1938" authorityName="(Luhe, 1906) Meserve" authorityYear="1938" class="Monogenea" family="Capsalidae" genus="Benedenia" kingdom="Animalia" order="Capsalidea" pageId="5" pageNumber="6" phylum="Platyhelminthes" rank="species" species="macrocolpa">
<emphasis id="B91C7047D31DFFECFC4242019EB3574E" box="[938,1184,1333,1356]" italics="true" pageId="5" pageNumber="6">Benedenia macrocolpa</emphasis>
(
<bibRefCitation id="EFF9D1A4D31DFFECFB4742019F38574E" author="Luhe" box="[1199,1323,1333,1356]" pageId="5" pageNumber="29" refString="Luhe, M. (1906) Report on the trematode parasites from the marine fishes of Ceylon. Report to the Government of Ceylon on the Pearl Oyster Fisheries of the Gulf of Manaar, Supplementary Report, 5, 97 - 108." type="journal article" year="1906">Lühe, 1906</bibRefCitation>
)
<bibRefCitation id="EFF9D1A4D31DFFECFAD142019B13576E" author="Meserve" pageId="5" pageNumber="29" refString="Meserve, F. G. (1938) Some monogenetic trematodes from the Galapagos Islands and the neighbouring Pacific. Reports of the Allan Hancock Pacific Expeditions (1932 - 1937), 2, 31 - 89." type="journal article" year="1938">Meserve, 1938</bibRefCitation>
</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection id="C372FFDED31DFFECFF7F42AE9B1D5598" pageId="5" pageNumber="6" type="materials_examined">
<paragraph id="8BD7AC55D31DFFECFF7F42AE9F7757B6" blockId="5.[151,1437,1434,2020]" box="[151,1380,1434,1460]" pageId="5" pageNumber="6">
<emphasis id="B91C7047D31DFFECFF7F42AE9B0A57B6" bold="true" box="[151,281,1434,1460]" pageId="5" pageNumber="6">
<typeStatus id="54D312F7D31DFFECFF7F42AE9AC757B6" box="[151,212,1434,1460]" pageId="5" pageNumber="6">Type</typeStatus>
host:
</emphasis>
<taxonomicName id="4C68D7D6D31DFFECFEC942AF999E57B6" authority="Muller &amp; Henle" authorityName="Muller &amp; Henle" box="[289,909,1434,1460]" class="Elasmobranchii" family="Myliobatidae" genus="Rhinoptera" kingdom="Animalia" order="Myliobatiformes" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="javanica">
<emphasis id="B91C7047D31DFFECFEC942AF980257B6" box="[289,529,1435,1460]" italics="true" pageId="5" pageNumber="6">Rhinoptera javanica</emphasis>
Müller &amp; Henle (Myliobatidae)
</taxonomicName>
, the javanese cownose ray (
<bibRefCitation id="EFF9D1A4D31DFFECFB3B42AE9F4457B6" author="Luhe" box="[1235,1367,1434,1460]" pageId="5" pageNumber="29" refString="Luhe, M. (1906) Report on the trematode parasites from the marine fishes of Ceylon. Report to the Government of Ceylon on the Pearl Oyster Fisheries of the Gulf of Manaar, Supplementary Report, 5, 97 - 108." type="journal article" year="1906">Lühe 1906</bibRefCitation>
).
</paragraph>
<paragraph id="8BD7AC55D31DFFECFF2E428B991557D8" blockId="5.[151,1437,1434,2020]" box="[198,774,1471,1498]" pageId="5" pageNumber="6">
<emphasis id="B91C7047D31DFFECFF2E428B9B7C57DB" bold="true" box="[198,367,1471,1497]" pageId="5" pageNumber="6">
<typeStatus id="54D312F7D31DFFECFF2E428B9B1757DB" box="[198,260,1471,1497]" pageId="5" pageNumber="6">Type</typeStatus>
locality:
</emphasis>
Kalpitiya,
<collectingCountry id="F37FECC5D31DFFECFE1B42F4987657D8" box="[499,613,1472,1498]" name="Sri Lanka" pageId="5" pageNumber="6">Sri Lanka</collectingCountry>
(
<bibRefCitation id="EFF9D1A4D31DFFECFD9D42F498EA57D8" author="Luhe" box="[629,761,1472,1498]" pageId="5" pageNumber="29" refString="Luhe, M. (1906) Report on the trematode parasites from the marine fishes of Ceylon. Report to the Government of Ceylon on the Pearl Oyster Fisheries of the Gulf of Manaar, Supplementary Report, 5, 97 - 108." type="journal article" year="1906">Lühe 1906</bibRefCitation>
).
</paragraph>
<paragraph id="8BD7AC55D31DFFECFF2E42D19B1D5598" blockId="5.[151,1437,1434,2020]" pageId="5" pageNumber="6">
<emphasis id="B91C7047D31DFFECFF2E42D198C857FD" bold="true" box="[198,731,1509,1535]" pageId="5" pageNumber="6">Other published host and locality records:</emphasis>
<taxonomicName id="4C68D7D6D31DFFECFD0B42D199E957FD" authority="Dutch Bay" authorityName="Dutch Bay" box="[739,1018,1509,1535]" class="Elasmobranchii" family="Myliobatidae" genus="Rhinoptera" kingdom="Animalia" order="Myliobatiformes" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="javanica">
<emphasis id="B91C7047D31DFFECFD0B42D1997957FC" box="[739,874,1509,1534]" italics="true" pageId="5" pageNumber="6">R. javanica</emphasis>
, Dutch Bay
</taxonomicName>
,
<collectingCountry id="F37FECC5D31DFFECFBEF42D19E6957FD" box="[1031,1146,1509,1535]" name="Sri Lanka" pageId="5" pageNumber="6">Sri Lanka</collectingCountry>
(
<bibRefCitation id="EFF9D1A4D31DFFECFB6342D19F0357FD" author="Luhe" box="[1163,1296,1509,1535]" pageId="5" pageNumber="29" refString="Luhe, M. (1906) Report on the trematode parasites from the marine fishes of Ceylon. Report to the Government of Ceylon on the Pearl Oyster Fisheries of the Gulf of Manaar, Supplementary Report, 5, 97 - 108." type="journal article" year="1906">Lühe 1906</bibRefCitation>
); captive
<taxonomicName id="4C68D7D6D31DFFECFA6B42D19AEE5426" class="Elasmobranchii" family="Myliobatidae" genus="Rhinoptera" kingdom="Animalia" order="Myliobatiformes" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="javanica">
<emphasis id="B91C7047D31DFFECFA6B42D19AEE5426" italics="true" pageId="5" pageNumber="6">R. javanica</emphasis>
</taxonomicName>
in an aquarium at Atlantis The Palm Hotel, Dubai,
<collectingCountry id="F37FECC5D31DFFECFC80413E9E7F5426" box="[872,1132,1546,1572]" name="United Arab Emirates" pageId="5" pageNumber="6">United Arab Emirates</collectingCountry>
but caught from the wild north of Dubai, Arabian Gulf (present study);
<taxonomicName id="4C68D7D6D31DFFECFD2C4104995E544B" box="[708,845,1584,1609]" class="Elasmobranchii" family="Myliobatidae" genus="Rhinoptera" kingdom="Animalia" order="Myliobatiformes" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="javanica">
<emphasis id="B91C7047D31DFFECFD2C4104995E544B" box="[708,845,1584,1609]" italics="true" pageId="5" pageNumber="6">R. javanica</emphasis>
</taxonomicName>
[but reevaluated and redetermined in the present study as
<taxonomicName id="4C68D7D6D31DFFECFEE441619819546D" authority="Ogilby" authorityName="Ogilby" box="[268,522,1621,1647]" class="Elasmobranchii" family="Myliobatidae" genus="Rhinoptera" kingdom="Animalia" order="Myliobatiformes" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="neglecta">
<emphasis id="B91C7047D31DFFECFEE441619B8C546C" box="[268,415,1621,1646]" italics="true" pageId="5" pageNumber="6">R. neglecta</emphasis>
Ogilby
</taxonomicName>
], Moreton Bay (
<geoCoordinate id="EE5CCA92D31DFFECFD374161995E546D" box="[735,845,1621,1647]" direction="south" orientation="latitude" pageId="5" pageNumber="6" precision="925" value="-27.416666">27°25’S</geoCoordinate>
,
<geoCoordinate id="EE5CCA92D31DFFECFCB4416199CE546D" box="[860,989,1621,1647]" direction="east" orientation="longitude" pageId="5" pageNumber="6" precision="925" value="153.33333">153°20’E</geoCoordinate>
), Queensland,
<collectingCountry id="F37FECC5D31DFFECFB4841619F0B546D" box="[1184,1304,1621,1647]" name="Australia" pageId="5" pageNumber="6">Australia</collectingCountry>
(
<bibRefCitation id="EFF9D1A4D31DFFECFAC241619B7A5496" author="Kearn" pageId="5" pageNumber="28" refString="Kearn, G. C. &amp; Whittington, I. D. (1991) Swimming in a sub-adult monogenean of the genus Entobdella. International Journal for Parasitology, 21, 739 - 741." type="journal article" year="1991">Kearn &amp; Whittington 1991</bibRefCitation>
,
<bibRefCitation id="EFF9D1A4D31DFFECFE9D414E9BA05496" author="Whittington" box="[373,435,1658,1684]" pageId="5" pageNumber="28" refString="Kearn, G. C. &amp; Whittington, I. D. (1992) Diversity of reproductive behaviour in platyhelminth parasites: insemination in some benedeniine (capsalid) monogeneans. Parasitology, 104, 489 - 496." type="journal article" year="1992">1992</bibRefCitation>
; present study
<emphasis id="B91C7047D31DFFECFDB2414E98775496" bold="true" box="[602,612,1658,1684]" pageId="5" pageNumber="6">)</emphasis>
; captive
<taxonomicName id="4C68D7D6D31DFFECFD27414F99405496" box="[719,851,1659,1684]" class="Elasmobranchii" family="Myliobatidae" genus="Rhinoptera" kingdom="Animalia" order="Myliobatiformes" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="neglecta">
<emphasis id="B91C7047D31DFFECFD27414F99405496" box="[719,851,1659,1684]" italics="true" pageId="5" pageNumber="6">R. neglecta</emphasis>
</taxonomicName>
at Reef HQ Aquarium, a public display aquarium in Townsville, Queensland,
<collectingCountry id="F37FECC5D31DFFECFE054194987354B8" box="[493,608,1696,1722]" name="Australia" pageId="5" pageNumber="6">Australia</collectingCountry>
, but hosts initially collected in a creek flowing into the Ross River, Townsville, Queensland,
<collectingCountry id="F37FECC5D31DFFECFE2C41F1982354DD" box="[452,560,1733,1759]" name="Australia" pageId="5" pageNumber="6">Australia</collectingCountry>
(present study);
<taxonomicName id="4C68D7D6D31DFFECFD1141F199B654DC" box="[761,933,1733,1759]" class="Elasmobranchii" family="Myliobatidae" genus="Rhinoptera" kingdom="Animalia" order="Myliobatiformes" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="neglecta">
<emphasis id="B91C7047D31DFFECFD1141F1990054DC" box="[761,787,1733,1758]" italics="true" pageId="5" pageNumber="6">R.</emphasis>
cf.
<emphasis id="B91C7047D31DFFECFCA941F199B654DC" box="[833,933,1733,1758]" italics="true" pageId="5" pageNumber="6">neglecta</emphasis>
</taxonomicName>
(autopsy numbers CMO 3-31, CMO3-48) and
<taxonomicName id="4C68D7D6D31DFFECFF2541DF9B485506" box="[205,347,1771,1796]" class="Elasmobranchii" family="Myliobatidae" genus="Rhinoptera" kingdom="Animalia" order="Myliobatiformes" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis id="B91C7047D31DFFECFF2541DF9B485506" box="[205,347,1771,1796]" italics="true" pageId="5" pageNumber="6">Rhinoptera</emphasis>
</taxonomicName>
sp. (autopsy number CMO3-43) off Weipa (
<geoCoordinate id="EE5CCA92D31DFFECFC7D41DE99E85506" box="[917,1019,1770,1796]" direction="south" orientation="latitude" pageId="5" pageNumber="6" precision="925" value="-12.666667">12°40'S</geoCoordinate>
,
<geoCoordinate id="EE5CCA92D31DFFECFBE241DE9E915506" box="[1034,1154,1770,1796]" direction="east" orientation="longitude" pageId="5" pageNumber="6" precision="925" value="141.86667">141°52'E</geoCoordinate>
), Gulf of Carpentaria, Queensland,
<collectingCountry id="F37FECC5D31DFFECFEDA40249B8C5528" box="[306,415,1808,1834]" name="Australia" pageId="5" pageNumber="6">Australia</collectingCountry>
(present study). Note that CMO3 numbers are unique autopsy identifiers assigned by collectors Professor J. Caira and Dr K. Jensen and can be used to access additional host information at a website dedicated to surveys of elasmobranchs for metazoan parasites at http://web2.uconn.edu/tapeworm/ hosts.php.
</paragraph>
</subSubSection>
<subSubSection id="C372FFDED31DFFE0FF2E40919E3954E7" lastPageId="9" lastPageNumber="10" pageId="5" pageNumber="6" type="description">
<paragraph id="8BD7AC55D31DFFECFF2E40919AFB55E6" blockId="5.[151,1437,1434,2020]" pageId="5" pageNumber="6">
<emphasis id="B91C7047D31DFFECFF2E40919B8055BD" bold="true" box="[198,403,1957,1983]" pageId="5" pageNumber="6">Site of infection:</emphasis>
Skin according to
<bibRefCitation id="EFF9D1A4D31DFFECFD984091991655BD" author="Luhe" box="[624,773,1957,1983]" pageId="5" pageNumber="29" refString="Luhe, M. (1906) Report on the trematode parasites from the marine fishes of Ceylon. Report to the Government of Ceylon on the Pearl Oyster Fisheries of the Gulf of Manaar, Supplementary Report, 5, 97 - 108." type="journal article" year="1906">Lühe (1906)</bibRefCitation>
; ventral skin (
<bibRefCitation id="EFF9D1A4D31DFFECFC4140919EE255BD" author="Kearn" box="[937,1265,1957,1983]" pageId="5" pageNumber="28" refString="Kearn, G. C. &amp; Whittington, I. D. (1991) Swimming in a sub-adult monogenean of the genus Entobdella. International Journal for Parasitology, 21, 739 - 741." type="journal article" year="1991">Kearn &amp; Whittington, 1991</bibRefCitation>
,
<bibRefCitation id="EFF9D1A4D31DFFECFB1640919F2855BD" author="Whittington" box="[1278,1339,1957,1983]" pageId="5" pageNumber="28" refString="Kearn, G. C. &amp; Whittington, I. D. (1992) Diversity of reproductive behaviour in platyhelminth parasites: insemination in some benedeniine (capsalid) monogeneans. Parasitology, 104, 489 - 496." type="journal article" year="1992">1992</bibRefCitation>
, present study).
</paragraph>
<caption id="DF17FCDDD31EFFEFFF7F41EF9B5F55F0" httpUri="https://zenodo.org/record/196224/files/figure.png" pageId="6" pageNumber="7" targetBox="[278,1303,216,1729]" targetPageId="6">
<paragraph id="8BD7AC55D31EFFEFFF7F41EF9B5F55F0" blockId="6.[151,1436,1755,2034]" pageId="6" pageNumber="7">
<emphasis id="B91C7047D31EFFEFFF7F41EF9B0C54F1" bold="true" box="[151,287,1755,1779]" pageId="6" pageNumber="7">FIGURE 1.</emphasis>
<taxonomicName id="4C68D7D6D31EFFEFFECE41EF998E54F0" authority="(Luhe, 1906) Yamaguti, 1963" authorityName="(Luhe, 1906) Yamaguti" authorityYear="1963" box="[294,925,1755,1778]" class="Monogenea" family="Capsalidae" genus="Benedeniella" kingdom="Animalia" order="Capsalidea" pageId="6" pageNumber="7" phylum="Platyhelminthes" rank="species" species="macrocolpa">
<emphasis id="B91C7047D31EFFEFFECE41EF985754F0" box="[294,580,1755,1778]" italics="true" pageId="6" pageNumber="7">Benedeniella macrocolpa</emphasis>
(Lühe, 1906) Yamaguti, 1963
</taxonomicName>
. Whole adult parasite, composite drawing, in ventral view based on AHC 29906 (specimen labeled no. 1) with some additions by hand. For clarity, extent of intestinal diverticula (entirely coextensive with the vitellarium) other than immediately posterior to pharynx and separate uterus parallel with terminal male genital duct are not shown and extensive convolutions of vas deferens are simplified (but are shown in full in Figure 6). Abbreviations: aao, anterior attachment organ; ah, anterior hamulus; as, accessory sclerite; c, cirrus; cp, common genital pore; cs, cirrus sac; dh, dorsal horn; e, excretory bladder; em, extrinsic muscle; g, germarium; gr, ventral groove; h, haptor; ho, hooklet; i, intestine; mv, marginal valve; n, longitudinal nerve trunk; o, ootype; p, pharynx; ph, posterior hamulus; rp, rounded papilla; t, testis; te, tendon; v, vagina; vi, vitelline follicle; vp, vaginal pore. Scale bar: 1 mm.
</paragraph>
</caption>
<caption id="DF17FCDDD31FFFEEFF7F402799035589" httpUri="https://zenodo.org/record/196225/files/figure.png" pageId="7" pageNumber="8" targetBox="[307,1294,216,1780]" targetPageId="7">
<paragraph id="8BD7AC55D31FFFEEFF7F402799035589" blockId="7.[151,1437,1811,1931]" pageId="7" pageNumber="8">
<emphasis id="B91C7047D31FFFEEFF7F40279B095529" bold="true" box="[151,282,1811,1835]" pageId="7" pageNumber="8">FIGURE 2.</emphasis>
<taxonomicName id="4C68D7D6D31FFFEEFEC94020996D5529" authority="(Luhe, 1906) Yamaguti, 1963" authorityName="(Luhe, 1906) Yamaguti" authorityYear="1963" box="[289,894,1812,1835]" class="Monogenea" family="Capsalidae" genus="Benedeniella" kingdom="Animalia" order="Capsalidea" pageId="7" pageNumber="8" phylum="Platyhelminthes" rank="species" species="macrocolpa">
<emphasis id="B91C7047D31FFFEEFEC9402098275529" box="[289,564,1812,1835]" italics="true" pageId="7" pageNumber="8">Benedeniella macrocolpa</emphasis>
(Lühe, 1906) Yamaguti, 1963
</taxonomicName>
. Haptoral sclerites of adult parasite.
<emphasis id="B91C7047D31FFFEEFAE340279F365529" bold="true" box="[1291,1317,1811,1835]" pageId="7" pageNumber="8">A.</emphasis>
Accessory sclerite.
<emphasis id="B91C7047D31FFFEEFF1240079B075549" bold="true" box="[250,276,1843,1867]" pageId="7" pageNumber="8">B.</emphasis>
Anterior hamulus.
<emphasis id="B91C7047D31FFFEEFE1C400798035549" bold="true" box="[500,528,1843,1867]" pageId="7" pageNumber="8">C.</emphasis>
Posterior hamulus.
<emphasis id="B91C7047D31FFFEEFD1F400799025549" bold="true" box="[759,785,1843,1867]" pageId="7" pageNumber="8">D.</emphasis>
Hooklet.
<emphasis id="B91C7047D31FFFEEFC6D400799B35549" bold="true" box="[901,928,1843,1867]" pageId="7" pageNumber="8">E.</emphasis>
Proximal ends of anterior hamulus (ah) and accessory sclerite (as) showing main tendon path (stippled) plus branch (solid) to haptor tissue (see also Figure 3A). Scale bars: A–B, 100 µm; C, 50 µm; D, 10 µm; E, 200 µm.
</paragraph>
</caption>
<paragraph id="8BD7AC55D31FFFEEFF2E40809E8855CC" blockId="7.[198,1436,1972,2036]" box="[198,1179,1972,1998]" pageId="7" pageNumber="8">
<emphasis id="B91C7047D31FFFEEFF2E40809B8A55CC" bold="true" box="[198,409,1972,1998]" pageId="7" pageNumber="8">Infection details:</emphasis>
unknown for specimens from
<taxonomicName id="4C68D7D6D31FFFEEFCE84080999655CF" box="[768,901,1972,1997]" class="Elasmobranchii" family="Myliobatidae" genus="Rhinoptera" kingdom="Animalia" order="Myliobatiformes" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="javanica">
<emphasis id="B91C7047D31FFFEEFCE84080999655CF" box="[768,901,1972,1997]" italics="true" pageId="7" pageNumber="8">R. javanica</emphasis>
</taxonomicName>
from the Arabian Gulf.
</paragraph>
<paragraph id="8BD7AC55D31FFFE1FF2E40EE9F0A52FE" blockId="7.[198,1436,1972,2036]" lastBlockId="8.[151,1436,152,476]" lastPageId="8" lastPageNumber="9" pageId="7" pageNumber="8">
In February/
<date id="FFD68A95D31FFFEEFEB440EE980255F6" box="[348,529,2010,2036]" pageId="7" pageNumber="8" value="1989-03-04">
March 1989,
<specimenCount id="9D6E67DCD31FFFEEFDEA40EE980255F6" box="[514,529,2010,2036]" pageId="7" pageNumber="8" type="generic">4</specimenCount>
</date>
specimens of
<taxonomicName id="4C68D7D6D31FFFEEFD2E40EE995D55F1" box="[710,846,2010,2035]" class="Elasmobranchii" family="Myliobatidae" genus="Rhinoptera" kingdom="Animalia" order="Myliobatiformes" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="neglecta">
<emphasis id="B91C7047D31FFFEEFD2E40EE995D55F1" box="[710,846,2010,2035]" italics="true" pageId="7" pageNumber="8">R. neglecta</emphasis>
</taxonomicName>
from Moreton Bay, Queensland,
<collectingCountry id="F37FECC5D31FFFEEFB0440EE9F4455F6" box="[1260,1367,2010,2036]" name="Australia" pageId="7" pageNumber="8">Australia</collectingCountry>
were examined (disc width range:
<quantity id="4C9001B0D310FFE1FE0447AC986052B0" box="[492,627,152,178]" metricMagnitude="-1" metricUnit="m" metricValue="7.15" metricValueMax="10.0" metricValueMin="4.3" pageId="8" pageNumber="9" unit="cm" value="71.5" valueMax="100.0" valueMin="43.0">43–100 cm</quantity>
; sex undetermined), 2 of which were infected (disc widths:
<quantity id="4C9001B0D310FFE1FADB47AC9F6F52B0" box="[1331,1404,152,178]" metricMagnitude="-1" metricUnit="m" metricValue="4.3" pageId="8" pageNumber="9" unit="cm" value="43.0">43 cm</quantity>
[6 parasites including
<specimenCount id="9D6E67DCD310FFE1FE964789981552D5" box="[382,518,189,215]" pageId="8" pageNumber="9" type="juvenile">2 juveniles</specimenCount>
],
<quantity id="4C9001B0D310FFE1FDF14789987052D5" box="[537,611,189,215]" metricMagnitude="-1" metricUnit="m" metricValue="4.8" pageId="8" pageNumber="9" unit="cm" value="48.0">48 cm</quantity>
[
<specimenCount id="9D6E67DCD310FFE1FD9D478998F852D5" box="[629,747,189,215]" pageId="8" pageNumber="9" type="juvenile">1 juvenile</specimenCount>
parasite]). Prevalence 50%; infection intensity, 1–6 (3.5); these rays were dead for nearly 2 h by time of dissection and had live parasites on the ventral skin.
</paragraph>
<paragraph id="8BD7AC55D310FFE1FF2D463C9B3B5390" blockId="8.[151,1436,152,476]" pageId="8" pageNumber="9">
In
<date id="FFD68A95D310FFE1FF0D463C9B805320" box="[229,403,264,290]" pageId="8" pageNumber="9" value="1991-03-02">
March 1991,
<specimenCount id="9D6E67DCD310FFE1FE6C463C9B805320" box="[388,403,264,290]" pageId="8" pageNumber="9" type="generic">2</specimenCount>
</date>
specimens of
<taxonomicName id="4C68D7D6D310FFE1FDD4463C98D25323" box="[572,705,264,289]" class="Elasmobranchii" family="Myliobatidae" genus="Rhinoptera" kingdom="Animalia" order="Myliobatiformes" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="neglecta">
<emphasis id="B91C7047D310FFE1FDD4463C98D25323" box="[572,705,264,289]" italics="true" pageId="8" pageNumber="9">R. neglecta</emphasis>
</taxonomicName>
from Moreton Bay, Queensland,
<collectingCountry id="F37FECC5D310FFE1FBA5463C9EAB5320" box="[1101,1208,264,290]" name="Australia" pageId="8" pageNumber="9">Australia</collectingCountry>
were both infected (disc widths:
<quantity id="4C9001B0D310FFE1FED346199B945345" box="[315,391,301,327]" metricMagnitude="-1" metricUnit="m" metricValue="5.2" pageId="8" pageNumber="9" unit="cm" value="52.0">52 cm</quantity>
[1 parasite],
<quantity id="4C9001B0D310FFE1FDC1461998675345" box="[553,628,301,327]" metricMagnitude="-1" metricUnit="m" metricValue="6.2" pageId="8" pageNumber="9" unit="cm" value="62.0">62 cm</quantity>
[4 parasites]; host sex undetermined). Prevalence 100%; infection intensity, 1–4 (2.5); these rays were dead for nearly 3 h by time of dissection and had living parasites on the ventral skin.
</paragraph>
<paragraph id="8BD7AC55D310FFE1FF2D46A99BB053DE" blockId="8.[151,1436,152,476]" pageId="8" pageNumber="9">
Infection details unknown for material from captive
<taxonomicName id="4C68D7D6D310FFE1FCA846A999DB53B4" box="[832,968,413,438]" class="Elasmobranchii" family="Myliobatidae" genus="Rhinoptera" kingdom="Animalia" order="Myliobatiformes" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="neglecta">
<emphasis id="B91C7047D310FFE1FCA846A999DB53B4" box="[832,968,413,438]" italics="true" pageId="8" pageNumber="9">R. neglecta</emphasis>
</taxonomicName>
from Reef HQ Aquarium, Townsville, Queensland,
<collectingCountry id="F37FECC5D310FFE1FED846F69B8C53DE" box="[304,415,450,476]" name="Australia" pageId="8" pageNumber="9">Australia</collectingCountry>
.
</paragraph>
<caption id="DF17FCDDD310FFE1FF7F4160990055E9" httpUri="https://zenodo.org/record/196226/files/figure.png" pageId="8" pageNumber="9" targetBox="[248,1337,531,1594]" targetPageId="8">
<paragraph id="8BD7AC55D310FFE1FF7F4160990055E9" blockId="8.[151,1437,1620,2027]" pageId="8" pageNumber="9">
<emphasis id="B91C7047D310FFE1FF7F41609B0A546E" bold="true" box="[151,281,1620,1644]" pageId="8" pageNumber="9">FIGURE 3.</emphasis>
Differential interference contrast photomicrographs of haptor and anterior attachment organs from stained, mounted specimens of
<taxonomicName id="4C68D7D6D310FFE1FE75414098235489" box="[413,560,1652,1675]" class="Monogenea" family="Capsalidae" genus="Benedeniella" kingdom="Animalia" order="Capsalidea" pageId="8" pageNumber="9" phylum="Platyhelminthes" rank="genus">
<emphasis id="B91C7047D310FFE1FE75414098235489" box="[413,560,1652,1675]" italics="true" pageId="8" pageNumber="9">Benedeniella</emphasis>
</taxonomicName>
species.
<emphasis id="B91C7047D310FFE1FD72414098A6548E" bold="true" box="[666,693,1652,1676]" pageId="8" pageNumber="9">A.</emphasis>
Accessory sclerite (as) and anterior (= proximal) part of anterior hamulus (ah) on one side of haptor of
<taxonomicName id="4C68D7D6D310FFE1FDD641A09E3854A9" authority="(Luhe, 1906) Yamaguti, 1963" authorityName="(Luhe, 1906) Yamaguti" authorityYear="1963" box="[574,1067,1684,1707]" class="Monogenea" family="Capsalidae" genus="Benedeniella" kingdom="Animalia" order="Capsalidea" pageId="8" pageNumber="9" phylum="Platyhelminthes" rank="species" species="macrocolpa">
<emphasis id="B91C7047D310FFE1FDD641A098CC54A9" box="[574,735,1684,1707]" italics="true" pageId="8" pageNumber="9">B. macrocolpa</emphasis>
(Lühe, 1906) Yamaguti, 1963
</taxonomicName>
showing route of tendon through proximal notch (pn) of accessory sclerite before attaching to proximal end of anterior hamulus with a branch (b) of the tendon apparently attaching to haptor tissue. AHC 29906 (specimen labeled no. 1).
<emphasis id="B91C7047D310FFE1FBF941E09E3A54EE" bold="true" box="[1041,1065,1748,1772]" pageId="8" pageNumber="9">B.</emphasis>
The left anterior attachment organ of
<taxonomicName id="4C68D7D6D310FFE1FF5D41C09B455509" box="[181,342,1780,1803]" class="Monogenea" family="Capsalidae" genus="Benedeniella" kingdom="Animalia" order="Capsalidea" pageId="8" pageNumber="9" phylum="Platyhelminthes" rank="species" species="macrocolpa">
<emphasis id="B91C7047D310FFE1FF5D41C09AD65509" box="[181,197,1780,1803]" italics="true" pageId="8" pageNumber="9">B</emphasis>
.
<emphasis id="B91C7047D310FFE1FF3C41C09B455509" box="[212,342,1780,1803]" italics="true" pageId="8" pageNumber="9">macrocolpa</emphasis>
</taxonomicName>
showing complete ventral grooves (gr) and some incomplete grooves (ig) anterolaterally. Note the rounded papillae (rp) at the anterior extremity of the parasite. AHC 29906 (specimen labeled no. 1).
<emphasis id="B91C7047D310FFE1FAEE40209F32552E" bold="true" box="[1286,1313,1812,1836]" pageId="8" pageNumber="9">C.</emphasis>
Accessory sclerite (as) and anterior (= proximal) part of anterior hamulus (ah) on one side of haptor of
<taxonomicName id="4C68D7D6D310FFE1FB7840009B905569" authority="(Hargis, 1955) Yamaguti, 1963" authorityName="(Hargis, 1955) Yamaguti" authorityYear="1963" class="Monogenea" family="Capsalidae" genus="Benedenia" kingdom="Animalia" order="Capsalidea" pageId="8" pageNumber="9" phylum="Platyhelminthes" rank="species" species="posterocolpa">
<emphasis id="B91C7047D310FFE1FB7840009F2F5549" box="[1168,1340,1844,1867]" italics="true" pageId="8" pageNumber="9">B. posterocolpa</emphasis>
(Hargis, 1955) Yamaguti, 1963
</taxonomicName>
showing route of tendon through proximal notch of accessory sclerite before attaching to proximal end of anterior hamulus with a branch (b) of the tendon apparently attaching to haptor tissue. AHC 29911 (specimen labeled no. 13).
<emphasis id="B91C7047D310FFE1FEB940A09B7D55AE" bold="true" box="[337,366,1940,1964]" pageId="8" pageNumber="9">D.</emphasis>
The left anterior attachment organ of
<taxonomicName id="4C68D7D6D310FFE1FCCB40A099C555A9" box="[803,982,1940,1963]" class="Monogenea" family="Capsalidae" genus="Benedenia" kingdom="Animalia" order="Capsalidea" pageId="8" pageNumber="9" phylum="Platyhelminthes" rank="species" species="posterocolpa">
<emphasis id="B91C7047D310FFE1FCCB40A0992055A9" box="[803,819,1940,1963]" italics="true" pageId="8" pageNumber="9">B</emphasis>
.
<emphasis id="B91C7047D310FFE1FCAB40A099C555A9" box="[835,982,1940,1963]" italics="true" pageId="8" pageNumber="9">posterocolpa</emphasis>
</taxonomicName>
showing ventral grooves (gr) and some incomplete grooves (ig) anterolaterally. Note the dorsal horn (dh) where the body and anterior attachment organ join. AHC 29911 (specimen labeled no. 13). Scale bars: 200 µm.
</paragraph>
</caption>
<caption id="DF17FCDDD311FFE0FF7F4264989F5405" httpUri="https://zenodo.org/record/196227/files/figure.png" pageId="9" pageNumber="10" targetBox="[531,1055,180,1334]" targetPageId="9">
<paragraph id="8BD7AC55D311FFE0FF7F4264989F5405" blockId="9.[151,1437,1360,1543]" pageId="9" pageNumber="10">
<emphasis id="B91C7047D311FFE0FF7F42649B08576A" bold="true" box="[151,283,1360,1384]" pageId="9" pageNumber="10">FIGURE 4.</emphasis>
Scanning electron micrographs of dorsal region of juvenile specimen of
<taxonomicName id="4C68D7D6D311FFE0FBD042659B9F5785" authority="(Luhe, 1906) Yamaguti, 1963" authorityName="(Luhe, 1906) Yamaguti" authorityYear="1963" class="Monogenea" family="Capsalidae" genus="Benedeniella" kingdom="Animalia" order="Capsalidea" pageId="9" pageNumber="10" phylum="Platyhelminthes" rank="species" species="macrocolpa">
<emphasis id="B91C7047D311FFE0FBD042659F5E576A" box="[1080,1357,1361,1384]" italics="true" pageId="9" pageNumber="10">Benedeniella macrocolpa</emphasis>
(Lühe, 1906) Yamaguti, 1963
</taxonomicName>
.
<emphasis id="B91C7047D311FFE0FE7F42449BA3578A" bold="true" box="[407,432,1392,1416]" pageId="9" pageNumber="10">A.</emphasis>
Anterior region of specimen showing small tapering papillae resembling horns (h) dorsal to junction of body and anterior attachment organs. The position of the common genital pore (c) and the excretory papillae (ep) are also indicated.
<emphasis id="B91C7047D311FFE0FE4842849BA957CA" bold="true" box="[416,442,1456,1480]" pageId="9" pageNumber="10">B.</emphasis>
Haptor and posterior region of body proper showing pair of conical structures (co) near peduncle. Note marginal valve (mv).
<emphasis id="B91C7047D311FFE0FDC642E4985457EA" bold="true" box="[558,583,1488,1512]" pageId="9" pageNumber="10">C.</emphasis>
Enlargement of Figure 4B showing pair of conical structures dorsal to haptor. Scale bars: A, 250 µm; B, 250 µm; C, 100 µm.
</paragraph>
</caption>
<paragraph id="8BD7AC55D311FFE0FF2D41029E855499" blockId="9.[151,1437,1590,2027]" pageId="9" pageNumber="10">
In
<date id="FFD68A95D311FFE0FF0C41029B695452" box="[228,378,1590,1616]" pageId="9" pageNumber="10" value="2003-06-02">
June 2003,
<specimenCount id="9D6E67DCD311FFE0FE8341029B695452" box="[363,378,1590,1616]" pageId="9" pageNumber="10" type="male">2</specimenCount>
</date>
male specimens of
<taxonomicName id="4C68D7D6D311FFE0FD8A41029918544D" box="[610,779,1590,1616]" class="Elasmobranchii" family="Myliobatidae" genus="Rhinoptera" kingdom="Animalia" order="Myliobatiformes" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="neglecta">
<emphasis id="B91C7047D311FFE0FD8A4102986F544D" box="[610,636,1590,1615]" italics="true" pageId="9" pageNumber="10">R.</emphasis>
cf.
<emphasis id="B91C7047D311FFE0FD4041029918544D" box="[680,779,1590,1615]" italics="true" pageId="9" pageNumber="10">neglecta</emphasis>
</taxonomicName>
(CMO3-31, CMO3-48) off Weipa, Gulf of Carpentaria, Queensland,
<collectingCountry id="F37FECC5D311FFE0FEDF416F9BB45477" box="[311,423,1627,1653]" name="Australia" pageId="9" pageNumber="10">Australia</collectingCountry>
were infected (disc widths: CMO
<quantity id="4C9001B0D311FFE0FCB9416F99E75477" box="[849,1012,1627,1653]" metricMagnitude="-1" metricUnit="m" metricValue="1.70655" metricValueMax="3.1131" metricValueMin="0.3" pageId="9" pageNumber="10" unit="cm" value="17.0655" valueMax="31.131" valueMin="3.0">3–31, 131 cm</quantity>
[4 parasites including
<specimenCount id="9D6E67DCD311FFE0FAE7416F9F825477" box="[1295,1425,1627,1653]" pageId="9" pageNumber="10" type="juvenile">1 juvenile</specimenCount>
], CMO
<quantity id="4C9001B0D311FFE0FF3541B59B695499" box="[221,378,1665,1691]" metricMagnitude="-1" metricUnit="m" metricValue="2.5567" metricValueMax="4.8134" metricValueMin="0.3" pageId="9" pageNumber="10" unit="cm" value="25.567" valueMax="48.134" valueMin="3.0">3–48, 134 cm</quantity>
[
<specimenCount id="9D6E67DCD311FFE0FE6241B59BC85499" box="[394,475,1665,1691]" pageId="9" pageNumber="10" type="adult">5 adult</specimenCount>
parasites]). Prevalence 100%; infection intensity 4–5 (4.5).
</paragraph>
<paragraph id="8BD7AC55D311FFE0FF2D41929E3954E7" blockId="9.[151,1437,1590,2027]" pageId="9" pageNumber="10">
In
<date id="FFD68A95D311FFE0FF0C41929B6B54C2" box="[228,376,1702,1728]" pageId="9" pageNumber="10" value="2003-06-01">
June 2003,
<specimenCount id="9D6E67DCD311FFE0FE8141929B6B54C2" box="[361,376,1702,1728]" pageId="9" pageNumber="10" type="male">1</specimenCount>
</date>
male specimen of
<taxonomicName id="4C68D7D6D311FFE0FDBA419298C554BD" box="[594,726,1702,1727]" class="Elasmobranchii" family="Myliobatidae" genus="Rhinoptera" kingdom="Animalia" order="Myliobatiformes" pageId="9" pageNumber="10" phylum="Chordata" rank="genus">
<emphasis id="B91C7047D311FFE0FDBA419298C554BD" box="[594,726,1702,1727]" italics="true" pageId="9" pageNumber="10">Rhinoptera</emphasis>
</taxonomicName>
sp. (CMO3-43) off Weipa, Gulf of Carpentaria, Queensland,
<collectingCountry id="F37FECC5D311FFE0FF7F41FF9B1154E7" box="[151,258,1739,1765]" name="Australia" pageId="9" pageNumber="10">Australia</collectingCountry>
was infected (disc width:
<quantity id="4C9001B0D311FFE0FDDF41FF989D54E7" box="[567,654,1739,1765]" metricMagnitude="0" metricUnit="m" metricValue="1.27" pageId="9" pageNumber="10" unit="cm" value="127.0">127 cm</quantity>
[8 parasites including
<specimenCount id="9D6E67DCD311FFE0FC7141FF9E0C54E7" box="[921,1055,1739,1765]" pageId="9" pageNumber="10" type="juvenile">3 juveniles</specimenCount>
].
</paragraph>
</subSubSection>
<subSubSection id="C372FFDED311FFE4FF2E41C49F115578" lastPageId="13" lastPageNumber="14" pageId="9" pageNumber="10" type="materials_examined">
<paragraph id="8BD7AC55D311FFE3FF2E41C4990F53B5" blockId="9.[151,1437,1590,2027]" lastBlockId="10.[151,1437,152,439]" lastPageId="10" lastPageNumber="11" pageId="9" pageNumber="10">
<emphasis id="B91C7047D311FFE0FF2E41C49BAF5508" bold="true" box="[198,444,1776,1802]" pageId="9" pageNumber="10">Material deposited:</emphasis>
<collectionCode id="ED793490D311FFE0FE2C41C5980F5509" box="[452,540,1777,1803]" pageId="9" pageNumber="10">BMNH</collectionCode>
2010.4.16.1–3 (
<specimenCount id="9D6E67DCD311FFE0FD3541C599625509" box="[733,881,1777,1803]" pageId="9" pageNumber="10" type="generic">3 specimens</specimenCount>
, 3 slides),
<collectionCode id="ED793490D311FFE0FC0741C59E575509" LSID="urn:lsid:biocol.org:col:34982" box="[1007,1092,1777,1803]" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34982" name="South Australia Museum" pageId="9" pageNumber="10">SAMA</collectionCode>
<collectionCode id="ED793490D311FFE0FBA441C59E995509" box="[1100,1162,1777,1803]" pageId="9" pageNumber="10">AHC</collectionCode>
29900 (
<specimenCount id="9D6E67DCD311FFE0FB0441C59F925509" box="[1260,1409,1777,1803]" pageId="9" pageNumber="10" type="generic">4 specimens</specimenCount>
, 4 slides),
<collectionCode id="ED793490D311FFE0FF1E40229B445532" box="[246,343,1814,1840]" httpUri="http://grbio.org/cool/ppqr-es1r" name="United States National Parasite Collection" pageId="9" pageNumber="10">USNPC</collectionCode>
No.
<date id="FFD68A95D311FFE0FE7C40229BFD5532" box="[404,494,1814,1840]" pageId="9" pageNumber="10" value="09-10-30">103009</date>
–
<date id="FFD68A95D311FFE0FE16402298455532" box="[510,598,1814,1840]" pageId="9" pageNumber="10" value="11-10-30">103011</date>
(
<specimenCount id="9D6E67DCD311FFE0FD81402299125532" box="[617,769,1814,1840]" pageId="9" pageNumber="10" type="generic">3 specimens</specimenCount>
, 3 slides) ex captive specimens of
<taxonomicName id="4C68D7D6D311FFE0FB4240229F26552D" box="[1194,1333,1814,1839]" class="Elasmobranchii" family="Myliobatidae" genus="Rhinoptera" kingdom="Animalia" order="Myliobatiformes" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="javanica">
<emphasis id="B91C7047D311FFE0FB4240229F26552D" box="[1194,1333,1814,1839]" italics="true" pageId="9" pageNumber="10">R. javanica</emphasis>
</taxonomicName>
from an aquarium at Atlantis The Palm Hotel, Dubai,
<collectingCountry id="F37FECC5D311FFE0FD0B400F99EF5557" box="[739,1020,1851,1877]" name="United Arab Emirates" pageId="9" pageNumber="10">United Arab Emirates</collectingCountry>
(original source,
<collectionCode id="ED793490D311FFE0FB0A400F9F395557" box="[1250,1322,1851,1877]" httpUri="http://grbio.org/cool/nvww-iigc" name="Division of Standardisation" pageId="9" pageNumber="10">DBV</collectionCode>
);
<collectionCode id="ED793490D311FFE0FAA9400F9F8E5557" box="[1345,1437,1851,1877]" pageId="9" pageNumber="10">BMNH</collectionCode>
2010.4.16.4–5 (
<specimenCount id="9D6E67DCD311FFE0FE81405598005579" box="[361,531,1889,1915]" pageId="9" pageNumber="10" type="generic">2 specimens</specimenCount>
, 2 slides),
<collectionCode id="ED793490D311FFE0FD4F405599105579" LSID="urn:lsid:biocol.org:col:34982" box="[679,771,1889,1915]" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34982" name="South Australia Museum" pageId="9" pageNumber="10">SAMA</collectionCode>
<collectionCode id="ED793490D311FFE0FCE44055995C5579" box="[780,847,1889,1915]" pageId="9" pageNumber="10">AHC</collectionCode>
29901 (
<specimenCount id="9D6E67DCD311FFE0FC5740559E7B5579" box="[959,1128,1889,1915]" pageId="9" pageNumber="10" type="generic">3 specimens</specimenCount>
, 3 slides),
<collectionCode id="ED793490D311FFE0FB1440559F705579" box="[1276,1379,1889,1915]" httpUri="http://grbio.org/cool/ppqr-es1r" name="United States National Parasite Collection" pageId="9" pageNumber="10">USNPC</collectionCode>
No.
<date id="FFD68A95D311FFE0FF7F40B29AFC55A2" box="[151,239,1926,1952]" pageId="9" pageNumber="10" value="12-10-30">103012</date>
–
<date id="FFD68A95D311FFE0FF1640B29B4655A2" box="[254,341,1926,1952]" pageId="9" pageNumber="10" value="13-10-30">103013</date>
(
<specimenCount id="9D6E67DCD311FFE0FE8E40B29BE955A2" box="[358,506,1926,1952]" pageId="9" pageNumber="10" type="generic">2 specimens</specimenCount>
, 2 slides) ex
<taxonomicName id="4C68D7D6D311FFE0FD7D40B2990A559D" box="[661,793,1926,1951]" class="Elasmobranchii" family="Myliobatidae" genus="Rhinoptera" kingdom="Animalia" order="Myliobatiformes" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="neglecta">
<emphasis id="B91C7047D311FFE0FD7D40B2990A559D" box="[661,793,1926,1951]" italics="true" pageId="9" pageNumber="10">R. neglecta</emphasis>
</taxonomicName>
collected in 1989,
<collectionCode id="ED793490D311FFE0FC1340B29E4355A2" LSID="urn:lsid:biocol.org:col:34982" box="[1019,1104,1926,1952]" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34982" name="South Australia Museum" pageId="9" pageNumber="10">SAMA</collectionCode>
<collectionCode id="ED793490D311FFE0FBB040B29E8555A2" box="[1112,1174,1926,1952]" pageId="9" pageNumber="10">AHC</collectionCode>
29902 (
<specimenCount id="9D6E67DCD311FFE0FB1340B29F9155A2" box="[1275,1410,1926,1952]" pageId="9" pageNumber="10" type="generic">1 specimen</specimenCount>
, 1 slide), 2 sectioned specimens (referred to by
<bibRefCitation id="EFF9D1A4D311FFE0FD42409F99F955C7" author="Kearn" box="[682,1002,1963,1989]" pageId="9" pageNumber="28" refString="Kearn, G. C. &amp; Whittington, I. D. (1992) Diversity of reproductive behaviour in platyhelminth parasites: insemination in some benedeniine (capsalid) monogeneans. Parasitology, 104, 489 - 496." type="journal article" year="1992">Kearn &amp; Whittington 1992</bibRefCitation>
but not deposited until now)
<collectionCode id="ED793490D311FFE0FAAF409F9F8F55C7" LSID="urn:lsid:biocol.org:col:34982" box="[1351,1436,1963,1989]" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34982" name="South Australia Museum" pageId="9" pageNumber="10">SAMA</collectionCode>
<collectionCode id="ED793490D311FFE0FF7F40E59AC655E9" box="[151,213,2001,2027]" pageId="9" pageNumber="10">AHC</collectionCode>
29903 (15 slides of oblique 6 µm sections),
<collectionCode id="ED793490D311FFE0FD0040E5992E55E9" LSID="urn:lsid:biocol.org:col:34982" box="[744,829,2001,2027]" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34982" name="South Australia Museum" pageId="9" pageNumber="10">SAMA</collectionCode>
<collectionCode id="ED793490D311FFE0FCAD40E5999055E9" box="[837,899,2001,2027]" pageId="9" pageNumber="10">AHC</collectionCode>
29904 (9 slides of oblique 6 µm sections) ex
<taxonomicName id="4C68D7D6D312FFE3FF7F47AC9B3152B3" box="[151,290,152,177]" class="Elasmobranchii" family="Myliobatidae" genus="Rhinoptera" kingdom="Animalia" order="Myliobatiformes" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="neglecta">
<emphasis id="B91C7047D312FFE3FF7F47AC9B3152B3" box="[151,290,152,177]" italics="true" pageId="10" pageNumber="11">R. neglecta</emphasis>
</taxonomicName>
collected in
<date id="FFD68A95D312FFE3FE2A47AC984A52B0" box="[450,601,152,178]" pageId="10" pageNumber="11" value="1991-03">March 1991</date>
, Moreton Bay (
<geoCoordinate id="EE5CCA92D312FFE3FCF347AC999152B0" box="[795,898,152,178]" direction="south" orientation="latitude" pageId="10" pageNumber="11" precision="925" value="-27.416666">27°25’S</geoCoordinate>
,
<geoCoordinate id="EE5CCA92D312FFE3FC7947AC9E1952B0" box="[913,1034,152,178]" direction="east" orientation="longitude" pageId="10" pageNumber="11" precision="925" value="153.33333">153°20’E</geoCoordinate>
), Queensland,
<collectingCountry id="F37FECC5D312FFE3FB2947AC9F2652B0" box="[1217,1333,152,178]" name="Australia" pageId="10" pageNumber="11">Australia</collectingCountry>
;
<collectionCode id="ED793490D312FFE3FAAA47AC9F8F52B0" box="[1346,1436,152,178]" pageId="10" pageNumber="11">BMNH</collectionCode>
2010.4.16.6-8 (
<specimenCount id="9D6E67DCD312FFE3FE8E4789981C52D5" box="[358,527,189,215]" pageId="10" pageNumber="11" type="generic">3 specimens</specimenCount>
, 3 slides),
<collectionCode id="ED793490D312FFE3FD4C4789991352D5" LSID="urn:lsid:biocol.org:col:34982" box="[676,768,189,215]" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34982" name="South Australia Museum" pageId="10" pageNumber="11">SAMA</collectionCode>
<collectionCode id="ED793490D312FFE3FCE24789995E52D5" box="[778,845,189,215]" pageId="10" pageNumber="11">AHC</collectionCode>
29905 (
<specimenCount id="9D6E67DCD312FFE3FC5547899E7452D5" box="[957,1127,189,215]" pageId="10" pageNumber="11" type="generic">4 specimens</specimenCount>
, 4 slides),
<collectionCode id="ED793490D312FFE3FB1447899F7052D5" box="[1276,1379,189,215]" httpUri="http://grbio.org/cool/ppqr-es1r" name="United States National Parasite Collection" pageId="10" pageNumber="11">USNPC</collectionCode>
No.
<date id="FFD68A95D312FFE3FF7F47D69AE152FE" box="[151,242,226,252]" pageId="10" pageNumber="11" value="14-10-30">103014</date>
–
<date id="FFD68A95D312FFE3FEEA47D69B4E52FE" box="[258,349,226,252]" pageId="10" pageNumber="11" value="16-10-30">103016</date>
(
<specimenCount id="9D6E67DCD312FFE3FE9847D6981852FE" box="[368,523,226,252]" pageId="10" pageNumber="11" type="generic">3 specimens</specimenCount>
, 3 slides) ex
<taxonomicName id="4C68D7D6D312FFE3FD4747D6992952F9" box="[687,826,226,251]" class="Elasmobranchii" family="Myliobatidae" genus="Rhinoptera" kingdom="Animalia" order="Myliobatiformes" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="neglecta">
<emphasis id="B91C7047D312FFE3FD4747D6992952F9" box="[687,826,226,251]" italics="true" pageId="10" pageNumber="11">R. neglecta</emphasis>
</taxonomicName>
ex Reef HQ Aquarium, Townsville, Queensland,
<collectingCountry id="F37FECC5D312FFE3FF7F463C9B155320" box="[151,262,264,290]" name="Australia" pageId="10" pageNumber="11">Australia</collectingCountry>
;
<collectionCode id="ED793490D312FFE3FEF9463C9B7A5320" box="[273,361,264,290]" pageId="10" pageNumber="11">BMNH</collectionCode>
2010.4.16.9 (
<specimenCount id="9D6E67DCD312FFE3FDE6463C98805320" box="[526,659,264,290]" pageId="10" pageNumber="11" type="generic">1 specimen</specimenCount>
, 1 slide),
<collectionCode id="ED793490D312FFE3FCEB463C994B5320" LSID="urn:lsid:biocol.org:col:34982" box="[771,856,264,290]" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34982" name="South Australia Museum" pageId="10" pageNumber="11">SAMA</collectionCode>
<collectionCode id="ED793490D312FFE3FCB7463C998E5320" box="[863,925,264,290]" pageId="10" pageNumber="11">AHC</collectionCode>
29906 (
<specimenCount id="9D6E67DCD312FFE3FC15463C9E835320" box="[1021,1168,264,290]" pageId="10" pageNumber="11" type="generic">2 specimens</specimenCount>
, 2 slides),
<collectionCode id="ED793490D312FFE3FAE3463C9F795320" box="[1291,1386,264,290]" httpUri="http://grbio.org/cool/ppqr-es1r" name="United States National Parasite Collection" pageId="10" pageNumber="11">USNPC</collectionCode>
No.
<date id="FFD68A95D312FFE3FF7F46199AE15345" box="[151,242,301,327]" pageId="10" pageNumber="11" value="17-10-30">103017</date>
(
<specimenCount id="9D6E67DCD312FFE3FEEE46199B835345" box="[262,400,301,327]" pageId="10" pageNumber="11" type="generic">1 specimen</specimenCount>
) ex
<taxonomicName id="4C68D7D6D312FFE3FE2E461998645344" box="[454,631,301,327]" class="Elasmobranchii" family="Myliobatidae" genus="Rhinoptera" kingdom="Animalia" order="Myliobatiformes" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="neglecta">
<emphasis id="B91C7047D312FFE3FE2E46199BF35344" box="[454,480,301,326]" italics="true" pageId="10" pageNumber="11">R.</emphasis>
cf.
<emphasis id="B91C7047D312FFE3FDF8461998645344" box="[528,631,301,326]" italics="true" pageId="10" pageNumber="11">neglecta</emphasis>
</taxonomicName>
(CMO 3-31) off Weipa (
<geoCoordinate id="EE5CCA92D312FFE3FC4346199E1E5345" box="[939,1037,301,327]" direction="south" orientation="latitude" pageId="10" pageNumber="11" precision="925" value="-12.666667">12°40'S</geoCoordinate>
,
<geoCoordinate id="EE5CCA92D312FFE3FBF346199E9D5345" box="[1051,1166,301,327]" direction="east" orientation="longitude" pageId="10" pageNumber="11" precision="925" value="141.86667">141°52'E</geoCoordinate>
), Gulf of Carpentaria, Queensland,
<collectingCountry id="F37FECC5D312FFE3FED846669B8C536E" box="[304,415,338,364]" name="Australia" pageId="10" pageNumber="11">Australia</collectingCountry>
;
<collectionCode id="ED793490D312FFE3FE4346669810536E" box="[427,515,338,364]" pageId="10" pageNumber="11">BMNH</collectionCode>
2010.4.16.10–11 (
<specimenCount id="9D6E67DCD312FFE3FD0846669967536E" box="[736,884,338,364]" pageId="10" pageNumber="11" type="generic">2 specimens</specimenCount>
, 2 slides),
<collectionCode id="ED793490D312FFE3FC1946669E55536E" LSID="urn:lsid:biocol.org:col:34982" box="[1009,1094,338,364]" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34982" name="South Australia Museum" pageId="10" pageNumber="11">SAMA</collectionCode>
<collectionCode id="ED793490D312FFE3FBA646669E9F536E" box="[1102,1164,338,364]" pageId="10" pageNumber="11">AHC</collectionCode>
29907 (
<specimenCount id="9D6E67DCD312FFE3FB0646669F92536E" box="[1262,1409,338,364]" pageId="10" pageNumber="11" type="generic">3 specimens</specimenCount>
, 3 slides),
<collectionCode id="ED793490D312FFE3FF11464C9B485390" box="[249,347,376,402]" httpUri="http://grbio.org/cool/ppqr-es1r" name="United States National Parasite Collection" pageId="10" pageNumber="11">USNPC</collectionCode>
No.
<date id="FFD68A95D312FFE3FE72464C9BE55390" box="[410,502,376,402]" pageId="10" pageNumber="11" value="18-10-30">103018</date>
(
<specimenCount id="9D6E67DCD312FFE3FDE0464C98B55390" box="[520,678,376,402]" pageId="10" pageNumber="11" type="generic">2 specimens</specimenCount>
, 2 slides) ex
<taxonomicName id="4C68D7D6D312FFE3FCA3464C99C55393" box="[843,982,376,401]" class="Elasmobranchii" family="Myliobatidae" genus="Rhinoptera" kingdom="Animalia" order="Myliobatiformes" pageId="10" pageNumber="11" phylum="Chordata" rank="genus">
<emphasis id="B91C7047D312FFE3FCA3464C99C55393" box="[843,982,376,401]" italics="true" pageId="10" pageNumber="11">Rhinoptera</emphasis>
</taxonomicName>
sp. (CMO3-43) off Weipa (
<geoCoordinate id="EE5CCA92D312FFE3FADB464C9F845390" box="[1331,1431,376,402]" direction="south" orientation="latitude" pageId="10" pageNumber="11" precision="925" value="-12.666667">12°40'S</geoCoordinate>
,
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), Gulf of Carpentaria, Queensland,
<collectingCountry id="F37FECC5D312FFE3FD4046A9990453B5" box="[680,791,413,439]" name="Australia" pageId="10" pageNumber="11">Australia</collectingCountry>
.
</paragraph>
<caption id="DF17FCDDD312FFE3FF7F42119E8C579F" httpUri="https://zenodo.org/record/196228/files/figure.png" pageId="10" pageNumber="11" targetBox="[250,1335,504,1289]" targetPageId="10">
<paragraph id="8BD7AC55D312FFE3FF7F42119E8C579F" blockId="10.[151,1437,1317,1437]" pageId="10" pageNumber="11">
<emphasis id="B91C7047D312FFE3FF7F42119B32573F" bold="true" box="[151,289,1317,1341]" pageId="10" pageNumber="11">FIGURE 5.</emphasis>
Diagrammatic outlines of adult
<taxonomicName id="4C68D7D6D312FFE3FD7142129F09573F" authority="(Luhe, 1906) Yamaguti, 1963" authorityName="(Luhe, 1906) Yamaguti" authorityYear="1963" box="[665,1306,1318,1341]" class="Monogenea" family="Capsalidae" genus="Benedeniella" kingdom="Animalia" order="Capsalidea" pageId="10" pageNumber="11" phylum="Platyhelminthes" rank="species" species="macrocolpa">
<emphasis id="B91C7047D312FFE3FD71421299AE573F" box="[665,957,1318,1341]" italics="true" pageId="10" pageNumber="11">Benedeniella macrocolpa</emphasis>
(Lühe, 1906) Yamaguti, 1963
</taxonomicName>
<emphasis id="B91C7047D312FFE3FACA42119F5A573F" bold="true" box="[1314,1353,1317,1341]" pageId="10" pageNumber="11">(A)</emphasis>
and
<taxonomicName id="4C68D7D6D312FFE3FA6A42129893575F" authority="(Hargis, 1955) Yamaguti, 1963" authorityName="(Hargis, 1955) Yamaguti" authorityYear="1963" class="Monogenea" family="Capsalidae" genus="Benedenia" kingdom="Animalia" order="Capsalidea" pageId="10" pageNumber="11" phylum="Platyhelminthes" rank="species" species="posterocolpa">
<emphasis id="B91C7047D312FFE3FA6A42129B37575F" italics="true" pageId="10" pageNumber="11">B. posterocolpa</emphasis>
(Hargis, 1955) Yamaguti, 1963
</taxonomicName>
<emphasis id="B91C7047D312FFE3FD6F427198B9575F" bold="true" box="[647,682,1349,1373]" pageId="10" pageNumber="11">(B)</emphasis>
in dorsal view showing relative positions of paired anterior tapering horns (dh), common genital papilla (cgp; absent in
<taxonomicName id="4C68D7D6D312FFE3FD2F42529961577F" box="[711,882,1382,1405]" class="Monogenea" family="Capsalidae" genus="Benedenia" kingdom="Animalia" order="Capsalidea" pageId="10" pageNumber="11" phylum="Platyhelminthes" rank="species" species="posterocolpa">
<emphasis id="B91C7047D312FFE3FD2F42529961577F" box="[711,882,1382,1405]" italics="true" pageId="10" pageNumber="11">B. posterocolpa</emphasis>
</taxonomicName>
), paired excretory papillae (ep) and paired conical structures (co). Drawn from several wholemounted specimens viewed dorsally. Scale bar: 4 mm.
</paragraph>
</caption>
<paragraph id="8BD7AC55D312FFE2FF2E42FF99E153B5" blockId="10.[151,1437,1483,2032]" lastBlockId="11.[151,1437,152,2007]" lastPageId="11" lastPageNumber="12" pageId="10" pageNumber="11">
<emphasis id="B91C7047D312FFE3FF2E42FF9B6D57E7" bold="true" box="[198,382,1483,1509]" pageId="10" pageNumber="11">Redescription:</emphasis>
Based on several live juvenile and adult specimens from Moreton Bay, Queensland, 20 flattened wholemounted adults, 15 flattened wholemounted juveniles, two series of oblique sections of two adults and one adult specimen processed for SEM from
<collectingCountry id="F37FECC5D312FFE3FCA5412299AD5432" box="[845,958,1558,1584]" name="Australia" pageId="10" pageNumber="11">Australia</collectingCountry>
and 10 wholemounted adults from the Arabian Gulf. Measurements from 20 flattened wholemounted specimens (all sexually mature) from
<collectingCountry id="F37FECC5D312FFE3FF7F41559B155479" box="[151,262,1633,1659]" name="Australia" pageId="10" pageNumber="11">Australia</collectingCountry>
, except hooklets measured in 10 flattened wholemounted juvenile specimens from
<collectingCountry id="F37FECC5D312FFE3FB3F41559F555479" box="[1239,1350,1633,1659]" name="Australia" pageId="10" pageNumber="11">Australia</collectingCountry>
; length of everted cirrus measured in four wholemounts from the Arabian Gulf only. Body broadly oval (
<figureCitation id="1353B0D0D312FFE3FAEB41B29F5B54A2" box="[1283,1352,1670,1696]" captionStart="FIGURE 1" captionStartId="6.[151,258,1755,1779]" captionTargetBox="[278,1303,216,1729]" captionTargetId="figure@6.[267,1321,197,1745]" captionTargetPageId="6" captionText="FIGURE 1. Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963. Whole adult parasite, composite drawing, in ventral view based on AHC 29906 (specimen labeled no. 1) with some additions by hand. For clarity, extent of intestinal diverticula (entirely coextensive with the vitellarium) other than immediately posterior to pharynx and separate uterus parallel with terminal male genital duct are not shown and extensive convolutions of vas deferens are simplified (but are shown in full in Figure 6). Abbreviations: aao, anterior attachment organ; ah, anterior hamulus; as, accessory sclerite; c, cirrus; cp, common genital pore; cs, cirrus sac; dh, dorsal horn; e, excretory bladder; em, extrinsic muscle; g, germarium; gr, ventral groove; h, haptor; ho, hooklet; i, intestine; mv, marginal valve; n, longitudinal nerve trunk; o, ootype; p, pharynx; ph, posterior hamulus; rp, rounded papilla; t, testis; te, tendon; v, vagina; vi, vitelline follicle; vp, vaginal pore. Scale bar: 1 mm." httpUri="https://zenodo.org/record/196224/files/figure.png" pageId="10" pageNumber="11">Fig. 1</figureCitation>
). Total length (including haptor) and maximum breadth (at level of testes): 4.13–10.67 (7.18, n = 20) x 2.5–6.54 (4.58, n = 20) mm. Haptor 1.29–2.89 (2.21, n = 20) x 1.32–2.78 (2.06, n = 20) mm; wider anteriorly, tapering posteriorly (
<figureCitation id="1353B0D0D312FFE3FEC341C29B615512" box="[299,370,1782,1808]" captionStart="FIGURE 1" captionStartId="6.[151,258,1755,1779]" captionTargetBox="[278,1303,216,1729]" captionTargetId="figure@6.[267,1321,197,1745]" captionTargetPageId="6" captionText="FIGURE 1. Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963. Whole adult parasite, composite drawing, in ventral view based on AHC 29906 (specimen labeled no. 1) with some additions by hand. For clarity, extent of intestinal diverticula (entirely coextensive with the vitellarium) other than immediately posterior to pharynx and separate uterus parallel with terminal male genital duct are not shown and extensive convolutions of vas deferens are simplified (but are shown in full in Figure 6). Abbreviations: aao, anterior attachment organ; ah, anterior hamulus; as, accessory sclerite; c, cirrus; cp, common genital pore; cs, cirrus sac; dh, dorsal horn; e, excretory bladder; em, extrinsic muscle; g, germarium; gr, ventral groove; h, haptor; ho, hooklet; i, intestine; mv, marginal valve; n, longitudinal nerve trunk; o, ootype; p, pharynx; ph, posterior hamulus; rp, rounded papilla; t, testis; te, tendon; v, vagina; vi, vitelline follicle; vp, vaginal pore. Scale bar: 1 mm." httpUri="https://zenodo.org/record/196224/files/figure.png" pageId="10" pageNumber="11">Fig. 1</figureCitation>
). Accessory sclerites large, 301–582 (459, n = 40) long, near haptor centre; shaped like spear-head; proximal end with notch (
<figureCitation id="1353B0D0D312FFE3FDB9402F98885537" box="[593,667,1819,1845]" captionStart="FIGURE 1" captionStartId="6.[151,258,1755,1779]" captionTargetBox="[278,1303,216,1729]" captionTargetId="figure@6.[267,1321,197,1745]" captionTargetPageId="6" captionText="FIGURE 1. Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963. Whole adult parasite, composite drawing, in ventral view based on AHC 29906 (specimen labeled no. 1) with some additions by hand. For clarity, extent of intestinal diverticula (entirely coextensive with the vitellarium) other than immediately posterior to pharynx and separate uterus parallel with terminal male genital duct are not shown and extensive convolutions of vas deferens are simplified (but are shown in full in Figure 6). Abbreviations: aao, anterior attachment organ; ah, anterior hamulus; as, accessory sclerite; c, cirrus; cp, common genital pore; cs, cirrus sac; dh, dorsal horn; e, excretory bladder; em, extrinsic muscle; g, germarium; gr, ventral groove; h, haptor; ho, hooklet; i, intestine; mv, marginal valve; n, longitudinal nerve trunk; o, ootype; p, pharynx; ph, posterior hamulus; rp, rounded papilla; t, testis; te, tendon; v, vagina; vi, vitelline follicle; vp, vaginal pore. Scale bar: 1 mm." httpUri="https://zenodo.org/record/196224/files/figure.png" pageId="10" pageNumber="11">Figs 1</figureCitation>
,
<figureCitation id="1353B0D0D312FFE3FD41402F98AB5537" box="[681,696,1819,1845]" captionStart="FIGURE 2" captionStartId="7.[151,255,1811,1835]" captionTargetBox="[307,1294,216,1780]" captionTargetId="figure@7.[289,1304,194,1795]" captionTargetPageId="7" captionText="FIGURE 2. Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963. Haptoral sclerites of adult parasite. A. Accessory sclerite. B. Anterior hamulus. C. Posterior hamulus. D. Hooklet. E. Proximal ends of anterior hamulus (ah) and accessory sclerite (as) showing main tendon path (stippled) plus branch (solid) to haptor tissue (see also Figure 3 A). Scale bars: A – B, 100 µm; C, 50 µm; D, 10 µm; E, 200 µm." httpUri="https://zenodo.org/record/196225/files/figure.png" pageId="10" pageNumber="11">2</figureCitation>
A, 3A); distal tips pointed, directed anteriorly (
<figureCitation id="1353B0D0D312FFE3FB34402F9F335537" box="[1244,1312,1819,1845]" captionStart="FIGURE 1" captionStartId="6.[151,258,1755,1779]" captionTargetBox="[278,1303,216,1729]" captionTargetId="figure@6.[267,1321,197,1745]" captionTargetPageId="6" captionText="FIGURE 1. Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963. Whole adult parasite, composite drawing, in ventral view based on AHC 29906 (specimen labeled no. 1) with some additions by hand. For clarity, extent of intestinal diverticula (entirely coextensive with the vitellarium) other than immediately posterior to pharynx and separate uterus parallel with terminal male genital duct are not shown and extensive convolutions of vas deferens are simplified (but are shown in full in Figure 6). Abbreviations: aao, anterior attachment organ; ah, anterior hamulus; as, accessory sclerite; c, cirrus; cp, common genital pore; cs, cirrus sac; dh, dorsal horn; e, excretory bladder; em, extrinsic muscle; g, germarium; gr, ventral groove; h, haptor; ho, hooklet; i, intestine; mv, marginal valve; n, longitudinal nerve trunk; o, ootype; p, pharynx; ph, posterior hamulus; rp, rounded papilla; t, testis; te, tendon; v, vagina; vi, vitelline follicle; vp, vaginal pore. Scale bar: 1 mm." httpUri="https://zenodo.org/record/196224/files/figure.png" pageId="10" pageNumber="11">Fig. 1</figureCitation>
). Anterior hamuli large, stout, strongly recurved distally, 373–839 (665, n = 40) long, with hooked distal tips (
<figureCitation id="1353B0D0D312FFE3FAD940759F6B5559" box="[1329,1400,1857,1883]" captionStart="FIGURE 2" captionStartId="7.[151,255,1811,1835]" captionTargetBox="[307,1294,216,1780]" captionTargetId="figure@7.[289,1304,194,1795]" captionTargetPageId="7" captionText="FIGURE 2. Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963. Haptoral sclerites of adult parasite. A. Accessory sclerite. B. Anterior hamulus. C. Posterior hamulus. D. Hooklet. E. Proximal ends of anterior hamulus (ah) and accessory sclerite (as) showing main tendon path (stippled) plus branch (solid) to haptor tissue (see also Figure 3 A). Scale bars: A – B, 100 µm; C, 50 µm; D, 10 µm; E, 200 µm." httpUri="https://zenodo.org/record/196225/files/figure.png" pageId="10" pageNumber="11">Fig. 2</figureCitation>
B). Hooked distal tips of one or both anterior hamuli broken off and missing in 13 mounted specimens from Townsville and Weipa, Queensland,
<collectingCountry id="F37FECC5D312FFE3FDA940BF98A355A7" box="[577,688,1931,1957]" name="Australia" pageId="10" pageNumber="11">Australia</collectingCountry>
, leaving blunt tips. Posterior hamuli relatively small, 53–84 (74, n = 33) long, slender with hooked distal tips (
<figureCitation id="1353B0D0D312FFE3FD2B4085991E55C9" box="[707,781,1969,1995]" captionStart="FIGURE 2" captionStartId="7.[151,255,1811,1835]" captionTargetBox="[307,1294,216,1780]" captionTargetId="figure@7.[289,1304,194,1795]" captionTargetPageId="7" captionText="FIGURE 2. Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963. Haptoral sclerites of adult parasite. A. Accessory sclerite. B. Anterior hamulus. C. Posterior hamulus. D. Hooklet. E. Proximal ends of anterior hamulus (ah) and accessory sclerite (as) showing main tendon path (stippled) plus branch (solid) to haptor tissue (see also Figure 3 A). Scale bars: A – B, 100 µm; C, 50 µm; D, 10 µm; E, 200 µm." httpUri="https://zenodo.org/record/196225/files/figure.png" pageId="10" pageNumber="11">Fig. 2</figureCitation>
C); position at haptor periphery indicated by marginal indentation (
<figureCitation id="1353B0D0D312FFE3FED840E29B6455F2" box="[304,375,2006,2032]" captionStart="FIGURE 1" captionStartId="6.[151,258,1755,1779]" captionTargetBox="[278,1303,216,1729]" captionTargetId="figure@6.[267,1321,197,1745]" captionTargetPageId="6" captionText="FIGURE 1. Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963. Whole adult parasite, composite drawing, in ventral view based on AHC 29906 (specimen labeled no. 1) with some additions by hand. For clarity, extent of intestinal diverticula (entirely coextensive with the vitellarium) other than immediately posterior to pharynx and separate uterus parallel with terminal male genital duct are not shown and extensive convolutions of vas deferens are simplified (but are shown in full in Figure 6). Abbreviations: aao, anterior attachment organ; ah, anterior hamulus; as, accessory sclerite; c, cirrus; cp, common genital pore; cs, cirrus sac; dh, dorsal horn; e, excretory bladder; em, extrinsic muscle; g, germarium; gr, ventral groove; h, haptor; ho, hooklet; i, intestine; mv, marginal valve; n, longitudinal nerve trunk; o, ootype; p, pharynx; ph, posterior hamulus; rp, rounded papilla; t, testis; te, tendon; v, vagina; vi, vitelline follicle; vp, vaginal pore. Scale bar: 1 mm." httpUri="https://zenodo.org/record/196224/files/figure.png" pageId="10" pageNumber="11">Fig. 1</figureCitation>
). Hooklets 11–15 (13, n = 45) long (
<figureCitation id="1353B0D0D312FFE3FCD340E2999655F2" box="[827,901,2006,2032]" captionStart="FIGURE 2" captionStartId="7.[151,255,1811,1835]" captionTargetBox="[307,1294,216,1780]" captionTargetId="figure@7.[289,1304,194,1795]" captionTargetPageId="7" captionText="FIGURE 2. Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963. Haptoral sclerites of adult parasite. A. Accessory sclerite. B. Anterior hamulus. C. Posterior hamulus. D. Hooklet. E. Proximal ends of anterior hamulus (ah) and accessory sclerite (as) showing main tendon path (stippled) plus branch (solid) to haptor tissue (see also Figure 3 A). Scale bars: A – B, 100 µm; C, 50 µm; D, 10 µm; E, 200 µm." httpUri="https://zenodo.org/record/196225/files/figure.png" pageId="10" pageNumber="11">Fig. 2</figureCitation>
D); positions on posterior and posterolateral borders of haptor indicated by marginal indentations (
<figureCitation id="1353B0D0D313FFE2FCCF47AC997D52B0" box="[807,878,152,178]" captionStart="FIGURE 1" captionStartId="6.[151,258,1755,1779]" captionTargetBox="[278,1303,216,1729]" captionTargetId="figure@6.[267,1321,197,1745]" captionTargetPageId="6" captionText="FIGURE 1. Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963. Whole adult parasite, composite drawing, in ventral view based on AHC 29906 (specimen labeled no. 1) with some additions by hand. For clarity, extent of intestinal diverticula (entirely coextensive with the vitellarium) other than immediately posterior to pharynx and separate uterus parallel with terminal male genital duct are not shown and extensive convolutions of vas deferens are simplified (but are shown in full in Figure 6). Abbreviations: aao, anterior attachment organ; ah, anterior hamulus; as, accessory sclerite; c, cirrus; cp, common genital pore; cs, cirrus sac; dh, dorsal horn; e, excretory bladder; em, extrinsic muscle; g, germarium; gr, ventral groove; h, haptor; ho, hooklet; i, intestine; mv, marginal valve; n, longitudinal nerve trunk; o, ootype; p, pharynx; ph, posterior hamulus; rp, rounded papilla; t, testis; te, tendon; v, vagina; vi, vitelline follicle; vp, vaginal pore. Scale bar: 1 mm." httpUri="https://zenodo.org/record/196224/files/figure.png" pageId="11" pageNumber="12">Fig. 1</figureCitation>
). Tendons conspicuous, travelling posteriorly from extrinsic muscles associated with haptor on each side of body; passing through notch at proximal ends of accessory sclerites before dividing, with part attaching to proximal ends of anterior hamuli and part attaching to ventral haptor tissue (
<figureCitation id="1353B0D0D313FFE2FE51463C98165320" box="[441,517,264,290]" captionStart="FIGURE 1" captionStartId="6.[151,258,1755,1779]" captionTargetBox="[278,1303,216,1729]" captionTargetId="figure@6.[267,1321,197,1745]" captionTargetPageId="6" captionText="FIGURE 1. Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963. Whole adult parasite, composite drawing, in ventral view based on AHC 29906 (specimen labeled no. 1) with some additions by hand. For clarity, extent of intestinal diverticula (entirely coextensive with the vitellarium) other than immediately posterior to pharynx and separate uterus parallel with terminal male genital duct are not shown and extensive convolutions of vas deferens are simplified (but are shown in full in Figure 6). Abbreviations: aao, anterior attachment organ; ah, anterior hamulus; as, accessory sclerite; c, cirrus; cp, common genital pore; cs, cirrus sac; dh, dorsal horn; e, excretory bladder; em, extrinsic muscle; g, germarium; gr, ventral groove; h, haptor; ho, hooklet; i, intestine; mv, marginal valve; n, longitudinal nerve trunk; o, ootype; p, pharynx; ph, posterior hamulus; rp, rounded papilla; t, testis; te, tendon; v, vagina; vi, vitelline follicle; vp, vaginal pore. Scale bar: 1 mm." httpUri="https://zenodo.org/record/196224/files/figure.png" pageId="11" pageNumber="12">Figs 1</figureCitation>
,
<figureCitation id="1353B0D0D313FFE2FDFC463C98305320" box="[532,547,264,290]" captionStart="FIGURE 2" captionStartId="7.[151,255,1811,1835]" captionTargetBox="[307,1294,216,1780]" captionTargetId="figure@7.[289,1304,194,1795]" captionTargetPageId="7" captionText="FIGURE 2. Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963. Haptoral sclerites of adult parasite. A. Accessory sclerite. B. Anterior hamulus. C. Posterior hamulus. D. Hooklet. E. Proximal ends of anterior hamulus (ah) and accessory sclerite (as) showing main tendon path (stippled) plus branch (solid) to haptor tissue (see also Figure 3 A). Scale bars: A – B, 100 µm; C, 50 µm; D, 10 µm; E, 200 µm." httpUri="https://zenodo.org/record/196225/files/figure.png" pageId="11" pageNumber="12">2</figureCitation>
E, 3A). Marginal valve running anteriorly from position of hooklets IV on each side of haptor extending to anteriormost region of haptor; wider anteriorly; absence on posteriormost part of haptor (e.g.
<figureCitation id="1353B0D0D313FFE2FE9346669BAD536E" box="[379,446,338,364]" captionStart="FIGURE 1" captionStartId="6.[151,258,1755,1779]" captionTargetBox="[278,1303,216,1729]" captionTargetId="figure@6.[267,1321,197,1745]" captionTargetPageId="6" captionText="FIGURE 1. Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963. Whole adult parasite, composite drawing, in ventral view based on AHC 29906 (specimen labeled no. 1) with some additions by hand. For clarity, extent of intestinal diverticula (entirely coextensive with the vitellarium) other than immediately posterior to pharynx and separate uterus parallel with terminal male genital duct are not shown and extensive convolutions of vas deferens are simplified (but are shown in full in Figure 6). Abbreviations: aao, anterior attachment organ; ah, anterior hamulus; as, accessory sclerite; c, cirrus; cp, common genital pore; cs, cirrus sac; dh, dorsal horn; e, excretory bladder; em, extrinsic muscle; g, germarium; gr, ventral groove; h, haptor; ho, hooklet; i, intestine; mv, marginal valve; n, longitudinal nerve trunk; o, ootype; p, pharynx; ph, posterior hamulus; rp, rounded papilla; t, testis; te, tendon; v, vagina; vi, vitelline follicle; vp, vaginal pore. Scale bar: 1 mm." httpUri="https://zenodo.org/record/196224/files/figure.png" pageId="11" pageNumber="12">Fig. 1</figureCitation>
) due to narrow width and damage. Ventral haptor surface apapillate. Pair of conical structures dorsal to haptor near peduncle observed with SEM (
<figureCitation id="1353B0D0D313FFE2FC92464C99D25390" box="[890,961,376,402]" captionStart="FIGURE 4" captionStartId="9.[151,255,1360,1384]" captionTargetBox="[531,1055,180,1334]" captionTargetId="figure@9.[531,1056,176,1339]" captionTargetPageId="9" captionText="FIGURE 4. Scanning electron micrographs of dorsal region of juvenile specimen of Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963. A. Anterior region of specimen showing small tapering papillae resembling horns (h) dorsal to junction of body and anterior attachment organs. The position of the common genital pore (c) and the excretory papillae (ep) are also indicated. B. Haptor and posterior region of body proper showing pair of conical structures (co) near peduncle. Note marginal valve (mv). C. Enlargement of Figure 4 B showing pair of conical structures dorsal to haptor. Scale bars: A, 250 µm; B, 250 µm; C, 100 µm." httpUri="https://zenodo.org/record/196227/files/figure.png" pageId="11" pageNumber="12">Fig. 4</figureCitation>
B, C); less apparent in wholemounts but visible if specimens viewed with dorsal surface uppermost (e.g.
<figureCitation id="1353B0D0D313FFE2FC6F46A999DF53B5" box="[903,972,413,439]" captionStart="FIGURE 5" captionStartId="10.[151,259,1317,1341]" captionTargetBox="[250,1335,504,1289]" captionTargetId="figure@10.[243,1344,493,1297]" captionTargetPageId="10" captionText="FIGURE 5. Diagrammatic outlines of adult Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963 (A) and B. posterocolpa (Hargis, 1955) Yamaguti, 1963 (B) in dorsal view showing relative positions of paired anterior tapering horns (dh), common genital papilla (cgp; absent in B. posterocolpa), paired excretory papillae (ep) and paired conical structures (co). Drawn from several wholemounted specimens viewed dorsally. Scale bar: 4 mm." httpUri="https://zenodo.org/record/196228/files/figure.png" pageId="11" pageNumber="12">Fig. 5</figureCitation>
A).
</paragraph>
<paragraph id="8BD7AC55D313FFE2FF2E46F69AEE51C0" blockId="11.[151,1437,152,2007]" pageId="11" pageNumber="12">
Anterior attachment organs ventral, almost circular, saucer-like discs, 467–1,350 (872, n = 38) x 603–1,251 (868, n = 40); ventral anterolateral region bearing conspicuous, complete grooves (
<figureCitation id="1353B0D0D313FFE2FB1C46DC9F535000" box="[1268,1344,488,514]" captionStart="FIGURE 1" captionStartId="6.[151,258,1755,1779]" captionTargetBox="[278,1303,216,1729]" captionTargetId="figure@6.[267,1321,197,1745]" captionTargetPageId="6" captionText="FIGURE 1. Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963. Whole adult parasite, composite drawing, in ventral view based on AHC 29906 (specimen labeled no. 1) with some additions by hand. For clarity, extent of intestinal diverticula (entirely coextensive with the vitellarium) other than immediately posterior to pharynx and separate uterus parallel with terminal male genital duct are not shown and extensive convolutions of vas deferens are simplified (but are shown in full in Figure 6). Abbreviations: aao, anterior attachment organ; ah, anterior hamulus; as, accessory sclerite; c, cirrus; cp, common genital pore; cs, cirrus sac; dh, dorsal horn; e, excretory bladder; em, extrinsic muscle; g, germarium; gr, ventral groove; h, haptor; ho, hooklet; i, intestine; mv, marginal valve; n, longitudinal nerve trunk; o, ootype; p, pharynx; ph, posterior hamulus; rp, rounded papilla; t, testis; te, tendon; v, vagina; vi, vitelline follicle; vp, vaginal pore. Scale bar: 1 mm." httpUri="https://zenodo.org/record/196224/files/figure.png" pageId="11" pageNumber="12">Figs 1</figureCitation>
,
<figureCitation id="1353B0D0D313FFE2FAA746DC9F4E5000" box="[1359,1373,488,514]" captionStart="FIGURE 3" captionStartId="8.[151,254,1620,1644]" captionTargetBox="[248,1337,531,1594]" captionTargetId="figure@8.[247,1339,530,1595]" captionTargetPageId="8" captionText="FIGURE 3. Differential interference contrast photomicrographs of haptor and anterior attachment organs from stained, mounted specimens of Benedeniella species. A. Accessory sclerite (as) and anterior (= proximal) part of anterior hamulus (ah) on one side of haptor of B. macrocolpa (Lühe, 1906) Yamaguti, 1963 showing route of tendon through proximal notch (pn) of accessory sclerite before attaching to proximal end of anterior hamulus with a branch (b) of the tendon apparently attaching to haptor tissue. AHC 29906 (specimen labeled no. 1). B. The left anterior attachment organ of B. macrocolpa showing complete ventral grooves (gr) and some incomplete grooves (ig) anterolaterally. Note the rounded papillae (rp) at the anterior extremity of the parasite. AHC 29906 (specimen labeled no. 1). C. Accessory sclerite (as) and anterior (= proximal) part of anterior hamulus (ah) on one side of haptor of B. posterocolpa (Hargis, 1955) Yamaguti, 1963 showing route of tendon through proximal notch of accessory sclerite before attaching to proximal end of anterior hamulus with a branch (b) of the tendon apparently attaching to haptor tissue. AHC 29911 (specimen labeled no. 13). D. The left anterior attachment organ of B. posterocolpa showing ventral grooves (gr) and some incomplete grooves (ig) anterolaterally. Note the dorsal horn (dh) where the body and anterior attachment organ join. AHC 29911 (specimen labeled no. 13). Scale bars: 200 µm." httpUri="https://zenodo.org/record/196226/files/figure.png" pageId="11" pageNumber="12">3</figureCitation>
B) of variable number, 11–28 (18, n = 55). Number of grooves in juvenile individuals similar to adults. In addition to complete grooves, 0–9 (3, n = 28) incomplete grooves present, of different lengths, occupying positions between complete grooves (
<figureCitation id="1353B0D0D313FFE2FE0E456C983E5070" box="[486,557,600,626]" captionStart="FIGURE 3" captionStartId="8.[151,254,1620,1644]" captionTargetBox="[248,1337,531,1594]" captionTargetId="figure@8.[247,1339,530,1595]" captionTargetPageId="8" captionText="FIGURE 3. Differential interference contrast photomicrographs of haptor and anterior attachment organs from stained, mounted specimens of Benedeniella species. A. Accessory sclerite (as) and anterior (= proximal) part of anterior hamulus (ah) on one side of haptor of B. macrocolpa (Lühe, 1906) Yamaguti, 1963 showing route of tendon through proximal notch (pn) of accessory sclerite before attaching to proximal end of anterior hamulus with a branch (b) of the tendon apparently attaching to haptor tissue. AHC 29906 (specimen labeled no. 1). B. The left anterior attachment organ of B. macrocolpa showing complete ventral grooves (gr) and some incomplete grooves (ig) anterolaterally. Note the rounded papillae (rp) at the anterior extremity of the parasite. AHC 29906 (specimen labeled no. 1). C. Accessory sclerite (as) and anterior (= proximal) part of anterior hamulus (ah) on one side of haptor of B. posterocolpa (Hargis, 1955) Yamaguti, 1963 showing route of tendon through proximal notch of accessory sclerite before attaching to proximal end of anterior hamulus with a branch (b) of the tendon apparently attaching to haptor tissue. AHC 29911 (specimen labeled no. 13). D. The left anterior attachment organ of B. posterocolpa showing ventral grooves (gr) and some incomplete grooves (ig) anterolaterally. Note the dorsal horn (dh) where the body and anterior attachment organ join. AHC 29911 (specimen labeled no. 13). Scale bars: 200 µm." httpUri="https://zenodo.org/record/196226/files/figure.png" pageId="11" pageNumber="12">Fig. 3</figureCitation>
B). Pair of rounded papillae (sense organs?) at anterior extremity of body proper between anterior attachment organs (
<figureCitation id="1353B0D0D313FFE2FD72454998F65095" box="[666,741,637,663]" captionStart="FIGURE 1" captionStartId="6.[151,258,1755,1779]" captionTargetBox="[278,1303,216,1729]" captionTargetId="figure@6.[267,1321,197,1745]" captionTargetPageId="6" captionText="FIGURE 1. Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963. Whole adult parasite, composite drawing, in ventral view based on AHC 29906 (specimen labeled no. 1) with some additions by hand. For clarity, extent of intestinal diverticula (entirely coextensive with the vitellarium) other than immediately posterior to pharynx and separate uterus parallel with terminal male genital duct are not shown and extensive convolutions of vas deferens are simplified (but are shown in full in Figure 6). Abbreviations: aao, anterior attachment organ; ah, anterior hamulus; as, accessory sclerite; c, cirrus; cp, common genital pore; cs, cirrus sac; dh, dorsal horn; e, excretory bladder; em, extrinsic muscle; g, germarium; gr, ventral groove; h, haptor; ho, hooklet; i, intestine; mv, marginal valve; n, longitudinal nerve trunk; o, ootype; p, pharynx; ph, posterior hamulus; rp, rounded papilla; t, testis; te, tendon; v, vagina; vi, vitelline follicle; vp, vaginal pore. Scale bar: 1 mm." httpUri="https://zenodo.org/record/196224/files/figure.png" pageId="11" pageNumber="12">Figs 1</figureCitation>
,
<figureCitation id="1353B0D0D313FFE2FD1B454999115095" box="[755,770,637,663]" captionStart="FIGURE 3" captionStartId="8.[151,254,1620,1644]" captionTargetBox="[248,1337,531,1594]" captionTargetId="figure@8.[247,1339,530,1595]" captionTargetPageId="8" captionText="FIGURE 3. Differential interference contrast photomicrographs of haptor and anterior attachment organs from stained, mounted specimens of Benedeniella species. A. Accessory sclerite (as) and anterior (= proximal) part of anterior hamulus (ah) on one side of haptor of B. macrocolpa (Lühe, 1906) Yamaguti, 1963 showing route of tendon through proximal notch (pn) of accessory sclerite before attaching to proximal end of anterior hamulus with a branch (b) of the tendon apparently attaching to haptor tissue. AHC 29906 (specimen labeled no. 1). B. The left anterior attachment organ of B. macrocolpa showing complete ventral grooves (gr) and some incomplete grooves (ig) anterolaterally. Note the rounded papillae (rp) at the anterior extremity of the parasite. AHC 29906 (specimen labeled no. 1). C. Accessory sclerite (as) and anterior (= proximal) part of anterior hamulus (ah) on one side of haptor of B. posterocolpa (Hargis, 1955) Yamaguti, 1963 showing route of tendon through proximal notch of accessory sclerite before attaching to proximal end of anterior hamulus with a branch (b) of the tendon apparently attaching to haptor tissue. AHC 29911 (specimen labeled no. 13). D. The left anterior attachment organ of B. posterocolpa showing ventral grooves (gr) and some incomplete grooves (ig) anterolaterally. Note the dorsal horn (dh) where the body and anterior attachment organ join. AHC 29911 (specimen labeled no. 13). Scale bars: 200 µm." httpUri="https://zenodo.org/record/196226/files/figure.png" pageId="11" pageNumber="12">3</figureCitation>
B). Additional pair of tapering papillae resembling horns located dorsally, one on each anterior attachment organ near junction with body (
<figureCitation id="1353B0D0D313FFE2FBBA45969E8E50BE" box="[1106,1181,674,700]" captionStart="FIGURE 1" captionStartId="6.[151,258,1755,1779]" captionTargetBox="[278,1303,216,1729]" captionTargetId="figure@6.[267,1321,197,1745]" captionTargetPageId="6" captionText="FIGURE 1. Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963. Whole adult parasite, composite drawing, in ventral view based on AHC 29906 (specimen labeled no. 1) with some additions by hand. For clarity, extent of intestinal diverticula (entirely coextensive with the vitellarium) other than immediately posterior to pharynx and separate uterus parallel with terminal male genital duct are not shown and extensive convolutions of vas deferens are simplified (but are shown in full in Figure 6). Abbreviations: aao, anterior attachment organ; ah, anterior hamulus; as, accessory sclerite; c, cirrus; cp, common genital pore; cs, cirrus sac; dh, dorsal horn; e, excretory bladder; em, extrinsic muscle; g, germarium; gr, ventral groove; h, haptor; ho, hooklet; i, intestine; mv, marginal valve; n, longitudinal nerve trunk; o, ootype; p, pharynx; ph, posterior hamulus; rp, rounded papilla; t, testis; te, tendon; v, vagina; vi, vitelline follicle; vp, vaginal pore. Scale bar: 1 mm." httpUri="https://zenodo.org/record/196224/files/figure.png" pageId="11" pageNumber="12">Figs 1</figureCitation>
,
<figureCitation id="1353B0D0D313FFE2FB4445969EA850BE" box="[1196,1211,674,700]" captionStart="FIGURE 4" captionStartId="9.[151,255,1360,1384]" captionTargetBox="[531,1055,180,1334]" captionTargetId="figure@9.[531,1056,176,1339]" captionTargetPageId="9" captionText="FIGURE 4. Scanning electron micrographs of dorsal region of juvenile specimen of Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963. A. Anterior region of specimen showing small tapering papillae resembling horns (h) dorsal to junction of body and anterior attachment organs. The position of the common genital pore (c) and the excretory papillae (ep) are also indicated. B. Haptor and posterior region of body proper showing pair of conical structures (co) near peduncle. Note marginal valve (mv). C. Enlargement of Figure 4 B showing pair of conical structures dorsal to haptor. Scale bars: A, 250 µm; B, 250 µm; C, 100 µm." httpUri="https://zenodo.org/record/196227/files/figure.png" pageId="11" pageNumber="12">4</figureCitation>
A, 5A); affinity for carmine in stained specimens. Eyes two pairs, pigment-shielded, dorsal, anterior to pharynx. Pharynx conspicuous, 387–1,106 (717, n = 20) x 404–976 (693, n = 20); intrinsic gland cells numerous posteriorly, staining with Ehrlich's haematoxylin and carmine or conspicuously granular in unstained specimens. Intestinal caeca branching extensively in body proper, coextensive with and mostly obscured by vitellarium; unclear whether gut caeca confluent posteriorly. Excretory bladders mostly conspicuous, each opening dorsally at level of posterior region of pharynx on distinct excretory papilla (
<figureCitation id="1353B0D0D313FFE2FBF144B69E70519E" box="[1049,1123,898,924]" captionStart="FIGURE 1" captionStartId="6.[151,258,1755,1779]" captionTargetBox="[278,1303,216,1729]" captionTargetId="figure@6.[267,1321,197,1745]" captionTargetPageId="6" captionText="FIGURE 1. Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963. Whole adult parasite, composite drawing, in ventral view based on AHC 29906 (specimen labeled no. 1) with some additions by hand. For clarity, extent of intestinal diverticula (entirely coextensive with the vitellarium) other than immediately posterior to pharynx and separate uterus parallel with terminal male genital duct are not shown and extensive convolutions of vas deferens are simplified (but are shown in full in Figure 6). Abbreviations: aao, anterior attachment organ; ah, anterior hamulus; as, accessory sclerite; c, cirrus; cp, common genital pore; cs, cirrus sac; dh, dorsal horn; e, excretory bladder; em, extrinsic muscle; g, germarium; gr, ventral groove; h, haptor; ho, hooklet; i, intestine; mv, marginal valve; n, longitudinal nerve trunk; o, ootype; p, pharynx; ph, posterior hamulus; rp, rounded papilla; t, testis; te, tendon; v, vagina; vi, vitelline follicle; vp, vaginal pore. Scale bar: 1 mm." httpUri="https://zenodo.org/record/196224/files/figure.png" pageId="11" pageNumber="12">Figs 1</figureCitation>
,
<figureCitation id="1353B0D0D313FFE2FB9A44B69E92519E" box="[1138,1153,898,924]" captionStart="FIGURE 4" captionStartId="9.[151,255,1360,1384]" captionTargetBox="[531,1055,180,1334]" captionTargetId="figure@9.[531,1056,176,1339]" captionTargetPageId="9" captionText="FIGURE 4. Scanning electron micrographs of dorsal region of juvenile specimen of Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963. A. Anterior region of specimen showing small tapering papillae resembling horns (h) dorsal to junction of body and anterior attachment organs. The position of the common genital pore (c) and the excretory papillae (ep) are also indicated. B. Haptor and posterior region of body proper showing pair of conical structures (co) near peduncle. Note marginal valve (mv). C. Enlargement of Figure 4 B showing pair of conical structures dorsal to haptor. Scale bars: A, 250 µm; B, 250 µm; C, 100 µm." httpUri="https://zenodo.org/record/196227/files/figure.png" pageId="11" pageNumber="12">4</figureCitation>
A, 5A), with affinity for carmine.
</paragraph>
<paragraph id="8BD7AC55D313FFE2FF2E44F99828551E" blockId="11.[151,1437,152,2007]" pageId="11" pageNumber="12">
Testes ovoid, juxtaposed, 491–1,327 (972, n = 32) x 395–1,276 (894, n = 32); penetrated by many dorsoventral muscle bundles (
<figureCitation id="1353B0D0D313FFE2FE1044C6982D560E" box="[504,574,1010,1036]" captionStart="FIGURE 1" captionStartId="6.[151,258,1755,1779]" captionTargetBox="[278,1303,216,1729]" captionTargetId="figure@6.[267,1321,197,1745]" captionTargetPageId="6" captionText="FIGURE 1. Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963. Whole adult parasite, composite drawing, in ventral view based on AHC 29906 (specimen labeled no. 1) with some additions by hand. For clarity, extent of intestinal diverticula (entirely coextensive with the vitellarium) other than immediately posterior to pharynx and separate uterus parallel with terminal male genital duct are not shown and extensive convolutions of vas deferens are simplified (but are shown in full in Figure 6). Abbreviations: aao, anterior attachment organ; ah, anterior hamulus; as, accessory sclerite; c, cirrus; cp, common genital pore; cs, cirrus sac; dh, dorsal horn; e, excretory bladder; em, extrinsic muscle; g, germarium; gr, ventral groove; h, haptor; ho, hooklet; i, intestine; mv, marginal valve; n, longitudinal nerve trunk; o, ootype; p, pharynx; ph, posterior hamulus; rp, rounded papilla; t, testis; te, tendon; v, vagina; vi, vitelline follicle; vp, vaginal pore. Scale bar: 1 mm." httpUri="https://zenodo.org/record/196224/files/figure.png" pageId="11" pageNumber="12">Fig. 1</figureCitation>
). Glands of Goto absent in juvenile and adult specimens. Vasa efferentia short (not drawn) uniting immediately posterior to germarium. Vas deferens travelling to left of germarium, passing dorsal to transverse vitelline ducts and proximal region of vagina, then becoming intensely convoluted, occupying space within curve formed by sac of male copulatory organ (= cirrus) posterior to ootype (
<figureCitation id="1353B0D0D313FFE2FF1F43BC9B5056A0" box="[247,323,1160,1186]" captionStart="FIGURE 1" captionStartId="6.[151,258,1755,1779]" captionTargetBox="[278,1303,216,1729]" captionTargetId="figure@6.[267,1321,197,1745]" captionTargetPageId="6" captionText="FIGURE 1. Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963. Whole adult parasite, composite drawing, in ventral view based on AHC 29906 (specimen labeled no. 1) with some additions by hand. For clarity, extent of intestinal diverticula (entirely coextensive with the vitellarium) other than immediately posterior to pharynx and separate uterus parallel with terminal male genital duct are not shown and extensive convolutions of vas deferens are simplified (but are shown in full in Figure 6). Abbreviations: aao, anterior attachment organ; ah, anterior hamulus; as, accessory sclerite; c, cirrus; cp, common genital pore; cs, cirrus sac; dh, dorsal horn; e, excretory bladder; em, extrinsic muscle; g, germarium; gr, ventral groove; h, haptor; ho, hooklet; i, intestine; mv, marginal valve; n, longitudinal nerve trunk; o, ootype; p, pharynx; ph, posterior hamulus; rp, rounded papilla; t, testis; te, tendon; v, vagina; vi, vitelline follicle; vp, vaginal pore. Scale bar: 1 mm." httpUri="https://zenodo.org/record/196224/files/figure.png" pageId="11" pageNumber="12">Figs 1</figureCitation>
,
<figureCitation id="1353B0D0D313FFE2FEBA43BC9B7056A0" box="[338,355,1160,1186]" captionStart="FIGURE 6" captionStartId="12.[151,256,1736,1760]" captionTargetBox="[282,1310,205,1705]" captionTargetId="figure@12.[270,1326,194,1716]" captionTargetPageId="12" captionText="FIGURE 6. Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963. Anterolateral region of body (left side) of adult parasite. Composite drawing in ventral view based on AHC 29905 (specimen labeled no. 1) with some additions by hand. Note that positions of some ducts (e. g. transverse vitelline duct, vas deferens and uterus) are displaced slightly for clarity. Also, common genital papilla, common genital pore, excretory papilla and vaginal pore are located dorsally and are shown here as seen by transparency. Abbreviations: cgp, common genital papilla; cv, common vitelline duct; dv, distal part of vagina; ep, excretory papilla; ey, eye; fc, fertilisation chamber; ju, junction of uterus and terminal male genital duct; m, male accessory gland reservoir; mg, terminal male genital duct; og, ootype glands; ov, oviduct; ovo, ovovitelline duct; pv, proximal part of vagina; tvd, transverse vitelline duct; u, uterus; vd, vas deferens; vdc, vas deferens within cirrus sac; vr, vitelline reservoir. Other abbreviations as for Figure 1. Scale bar: 1 mm." httpUri="https://zenodo.org/record/196229/files/figure.png" pageId="11" pageNumber="12">6</figureCitation>
); breaching muscular wall of cirrus sac dorsally at anterior level of ootype chamber. Inside cirrus sac, vas deferens narrows, following winding path dorsally to proximal end, then returning ventrally to base of cirrus (
<figureCitation id="1353B0D0D313FFE2FEAF43E69B8356EE" box="[327,400,1234,1260]" captionStart="FIGURE 6" captionStartId="12.[151,256,1736,1760]" captionTargetBox="[282,1310,205,1705]" captionTargetId="figure@12.[270,1326,194,1716]" captionTargetPageId="12" captionText="FIGURE 6. Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963. Anterolateral region of body (left side) of adult parasite. Composite drawing in ventral view based on AHC 29905 (specimen labeled no. 1) with some additions by hand. Note that positions of some ducts (e. g. transverse vitelline duct, vas deferens and uterus) are displaced slightly for clarity. Also, common genital papilla, common genital pore, excretory papilla and vaginal pore are located dorsally and are shown here as seen by transparency. Abbreviations: cgp, common genital papilla; cv, common vitelline duct; dv, distal part of vagina; ep, excretory papilla; ey, eye; fc, fertilisation chamber; ju, junction of uterus and terminal male genital duct; m, male accessory gland reservoir; mg, terminal male genital duct; og, ootype glands; ov, oviduct; ovo, ovovitelline duct; pv, proximal part of vagina; tvd, transverse vitelline duct; u, uterus; vd, vas deferens; vdc, vas deferens within cirrus sac; vr, vitelline reservoir. Other abbreviations as for Figure 1. Scale bar: 1 mm." httpUri="https://zenodo.org/record/196229/files/figure.png" pageId="11" pageNumber="12">Fig. 6</figureCitation>
). No evidence for widening of vas deferens or presence of sperm-filled reservoir inside cirrus sac, but structure and arrangement difficult to determine, even in sections, due to complexity and convolutions within cirrus sac.
<collectingCountry id="F37FECC5D313FFE2FDE2422998555735" box="[522,582,1309,1335]" name="Malta" pageId="11" pageNumber="12">Male</collectingCountry>
accessory gland reservoir single, lying ventrally in cirrus sac; source of male accessory secretion not identified; no ducts observed penetrating proximal region of cirrus sac. Cirrus sac wall provided with muscle fibres (not shown); length and extent of curvature of cirrus sac increasing with increase in size of parasite; in juvenile and some small adult specimens, distal portion lying diagonally in body posterior to pharynx with proximal part curving medianly immediately anterior to germarium and/or testes (
<figureCitation id="1353B0D0D313FFE2FF0042EC9B3D57F0" box="[232,302,1496,1522]" captionStart="FIGURE 7" captionStartId="13.[151,256,1243,1267]" captionTargetBox="[207,1398,209,1213]" captionTargetId="figure@13.[188,1398,194,1221]" captionTargetPageId="13" captionText="FIGURE 7. Diagrammatic comparison of curvature of cirrus sac in example juvenile (A) and adult (B) Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963. A. Juvenile specimen (AHC 29905 (specimen labeled no. 9); total length (including haptor) = 3.37 mm; anterior hamulus lengths = 366 and 376 µm. B. Adult specimen (AHC 29905 (specimen labeled no. 8 )); total length (including haptor) = 8.1 mm; anterior hamulus lengths = 718 and 743 µm. Abbreviations as for Figure 1. Scale bars: A, 300 µm; B, 1 mm." httpUri="https://zenodo.org/record/196230/files/figure.png" pageId="11" pageNumber="12">Fig. 7</figureCitation>
A); in most large specimens, proximal region of cirrus sac may curve anteriorly, dorsal to ootype chamber, forming a complete loop enclosing common vitelline duct, oviduct, ovo-vitelline duct, ootype and convolutions of vas deferens (
<figureCitation id="1353B0D0D313FFE2FE1041169853543E" box="[504,576,1570,1596]" captionStart="FIGURE 7" captionStartId="13.[151,256,1243,1267]" captionTargetBox="[207,1398,209,1213]" captionTargetId="figure@13.[188,1398,194,1221]" captionTargetPageId="13" captionText="FIGURE 7. Diagrammatic comparison of curvature of cirrus sac in example juvenile (A) and adult (B) Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963. A. Juvenile specimen (AHC 29905 (specimen labeled no. 9); total length (including haptor) = 3.37 mm; anterior hamulus lengths = 366 and 376 µm. B. Adult specimen (AHC 29905 (specimen labeled no. 8 )); total length (including haptor) = 8.1 mm; anterior hamulus lengths = 718 and 743 µm. Abbreviations as for Figure 1. Scale bars: A, 300 µm; B, 1 mm." httpUri="https://zenodo.org/record/196230/files/figure.png" pageId="11" pageNumber="12">Fig. 7</figureCitation>
B). Distal duct from male accessory gland reservoir and distal part of vas deferens uniting near base of cirrus at anterior level of ootype chamber (
<figureCitation id="1353B0D0D313FFE2FC0C417C9E3D5460" box="[996,1070,1608,1634]" captionStart="FIGURE 1" captionStartId="6.[151,258,1755,1779]" captionTargetBox="[278,1303,216,1729]" captionTargetId="figure@6.[267,1321,197,1745]" captionTargetPageId="6" captionText="FIGURE 1. Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963. Whole adult parasite, composite drawing, in ventral view based on AHC 29906 (specimen labeled no. 1) with some additions by hand. For clarity, extent of intestinal diverticula (entirely coextensive with the vitellarium) other than immediately posterior to pharynx and separate uterus parallel with terminal male genital duct are not shown and extensive convolutions of vas deferens are simplified (but are shown in full in Figure 6). Abbreviations: aao, anterior attachment organ; ah, anterior hamulus; as, accessory sclerite; c, cirrus; cp, common genital pore; cs, cirrus sac; dh, dorsal horn; e, excretory bladder; em, extrinsic muscle; g, germarium; gr, ventral groove; h, haptor; ho, hooklet; i, intestine; mv, marginal valve; n, longitudinal nerve trunk; o, ootype; p, pharynx; ph, posterior hamulus; rp, rounded papilla; t, testis; te, tendon; v, vagina; vi, vitelline follicle; vp, vaginal pore. Scale bar: 1 mm." httpUri="https://zenodo.org/record/196224/files/figure.png" pageId="11" pageNumber="12">Figs 1</figureCitation>
,
<figureCitation id="1353B0D0D313FFE2FBD5417C9E5D5460" box="[1085,1102,1608,1634]" captionStart="FIGURE 6" captionStartId="12.[151,256,1736,1760]" captionTargetBox="[282,1310,205,1705]" captionTargetId="figure@12.[270,1326,194,1716]" captionTargetPageId="12" captionText="FIGURE 6. Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963. Anterolateral region of body (left side) of adult parasite. Composite drawing in ventral view based on AHC 29905 (specimen labeled no. 1) with some additions by hand. Note that positions of some ducts (e. g. transverse vitelline duct, vas deferens and uterus) are displaced slightly for clarity. Also, common genital papilla, common genital pore, excretory papilla and vaginal pore are located dorsally and are shown here as seen by transparency. Abbreviations: cgp, common genital papilla; cv, common vitelline duct; dv, distal part of vagina; ep, excretory papilla; ey, eye; fc, fertilisation chamber; ju, junction of uterus and terminal male genital duct; m, male accessory gland reservoir; mg, terminal male genital duct; og, ootype glands; ov, oviduct; ovo, ovovitelline duct; pv, proximal part of vagina; tvd, transverse vitelline duct; u, uterus; vd, vas deferens; vdc, vas deferens within cirrus sac; vr, vitelline reservoir. Other abbreviations as for Figure 1. Scale bar: 1 mm." httpUri="https://zenodo.org/record/196229/files/figure.png" pageId="11" pageNumber="12">6</figureCitation>
). Cirrus long, sinuous, often twisted on itself, running anterolaterally inside terminal male genital duct, ventral to separate uterus, opening submarginally, slightly dorsal, on left side of body via common genital pore on short genital papilla at a level anterior to pharynx close to posterior margin of left anterior attachment organ (
<figureCitation id="1353B0D0D313FFE2FBA2418C9E8554D0" box="[1098,1174,1720,1746]" captionStart="FIGURE 1" captionStartId="6.[151,258,1755,1779]" captionTargetBox="[278,1303,216,1729]" captionTargetId="figure@6.[267,1321,197,1745]" captionTargetPageId="6" captionText="FIGURE 1. Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963. Whole adult parasite, composite drawing, in ventral view based on AHC 29906 (specimen labeled no. 1) with some additions by hand. For clarity, extent of intestinal diverticula (entirely coextensive with the vitellarium) other than immediately posterior to pharynx and separate uterus parallel with terminal male genital duct are not shown and extensive convolutions of vas deferens are simplified (but are shown in full in Figure 6). Abbreviations: aao, anterior attachment organ; ah, anterior hamulus; as, accessory sclerite; c, cirrus; cp, common genital pore; cs, cirrus sac; dh, dorsal horn; e, excretory bladder; em, extrinsic muscle; g, germarium; gr, ventral groove; h, haptor; ho, hooklet; i, intestine; mv, marginal valve; n, longitudinal nerve trunk; o, ootype; p, pharynx; ph, posterior hamulus; rp, rounded papilla; t, testis; te, tendon; v, vagina; vi, vitelline follicle; vp, vaginal pore. Scale bar: 1 mm." httpUri="https://zenodo.org/record/196224/files/figure.png" pageId="11" pageNumber="12">Figs 1</figureCitation>
,
<figureCitation id="1353B0D0D313FFE2FB4D418C9EA454D0" box="[1189,1207,1720,1746]" captionStart="FIGURE 6" captionStartId="12.[151,256,1736,1760]" captionTargetBox="[282,1310,205,1705]" captionTargetId="figure@12.[270,1326,194,1716]" captionTargetPageId="12" captionText="FIGURE 6. Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963. Anterolateral region of body (left side) of adult parasite. Composite drawing in ventral view based on AHC 29905 (specimen labeled no. 1) with some additions by hand. Note that positions of some ducts (e. g. transverse vitelline duct, vas deferens and uterus) are displaced slightly for clarity. Also, common genital papilla, common genital pore, excretory papilla and vaginal pore are located dorsally and are shown here as seen by transparency. Abbreviations: cgp, common genital papilla; cv, common vitelline duct; dv, distal part of vagina; ep, excretory papilla; ey, eye; fc, fertilisation chamber; ju, junction of uterus and terminal male genital duct; m, male accessory gland reservoir; mg, terminal male genital duct; og, ootype glands; ov, oviduct; ovo, ovovitelline duct; pv, proximal part of vagina; tvd, transverse vitelline duct; u, uterus; vd, vas deferens; vdc, vas deferens within cirrus sac; vr, vitelline reservoir. Other abbreviations as for Figure 1. Scale bar: 1 mm." httpUri="https://zenodo.org/record/196229/files/figure.png" pageId="11" pageNumber="12">6</figureCitation>
). Cirrus capable of eversion outside animal to length of 2.6–4.1 (3.5, n = 4 measured along curves from wholemounted adult specimens from Arabian Gulf) mm.
</paragraph>
<paragraph id="8BD7AC55D313FFE4FF2E401C9F935428" blockId="11.[151,1437,152,2007]" lastBlockId="13.[151,1437,1440,2026]" lastPageId="13" lastPageNumber="14" pageId="11" pageNumber="12">
Germarium immediately pretesticular with extensive, centrally located fertilisation chamber ventrally at mid-line (
<figureCitation id="1353B0D0D313FFE2FEE740799B4F5565" box="[271,348,1869,1895]" captionStart="FIGURE 1" captionStartId="6.[151,258,1755,1779]" captionTargetBox="[278,1303,216,1729]" captionTargetId="figure@6.[267,1321,197,1745]" captionTargetPageId="6" captionText="FIGURE 1. Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963. Whole adult parasite, composite drawing, in ventral view based on AHC 29906 (specimen labeled no. 1) with some additions by hand. For clarity, extent of intestinal diverticula (entirely coextensive with the vitellarium) other than immediately posterior to pharynx and separate uterus parallel with terminal male genital duct are not shown and extensive convolutions of vas deferens are simplified (but are shown in full in Figure 6). Abbreviations: aao, anterior attachment organ; ah, anterior hamulus; as, accessory sclerite; c, cirrus; cp, common genital pore; cs, cirrus sac; dh, dorsal horn; e, excretory bladder; em, extrinsic muscle; g, germarium; gr, ventral groove; h, haptor; ho, hooklet; i, intestine; mv, marginal valve; n, longitudinal nerve trunk; o, ootype; p, pharynx; ph, posterior hamulus; rp, rounded papilla; t, testis; te, tendon; v, vagina; vi, vitelline follicle; vp, vaginal pore. Scale bar: 1 mm." httpUri="https://zenodo.org/record/196224/files/figure.png" pageId="11" pageNumber="12">Figs 1</figureCitation>
,
<figureCitation id="1353B0D0D313FFE2FE8540799B6D5565" box="[365,382,1869,1895]" captionStart="FIGURE 6" captionStartId="12.[151,256,1736,1760]" captionTargetBox="[282,1310,205,1705]" captionTargetId="figure@12.[270,1326,194,1716]" captionTargetPageId="12" captionText="FIGURE 6. Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963. Anterolateral region of body (left side) of adult parasite. Composite drawing in ventral view based on AHC 29905 (specimen labeled no. 1) with some additions by hand. Note that positions of some ducts (e. g. transverse vitelline duct, vas deferens and uterus) are displaced slightly for clarity. Also, common genital papilla, common genital pore, excretory papilla and vaginal pore are located dorsally and are shown here as seen by transparency. Abbreviations: cgp, common genital papilla; cv, common vitelline duct; dv, distal part of vagina; ep, excretory papilla; ey, eye; fc, fertilisation chamber; ju, junction of uterus and terminal male genital duct; m, male accessory gland reservoir; mg, terminal male genital duct; og, ootype glands; ov, oviduct; ovo, ovovitelline duct; pv, proximal part of vagina; tvd, transverse vitelline duct; u, uterus; vd, vas deferens; vdc, vas deferens within cirrus sac; vr, vitelline reservoir. Other abbreviations as for Figure 1. Scale bar: 1 mm." httpUri="https://zenodo.org/record/196229/files/figure.png" pageId="11" pageNumber="12">6</figureCitation>
), containing ripe oocytes. Oviduct long, narrowing immediately anterior to transverse vitelline duct, convoluted, travelling anteriorly, joining relatively long common vitelline duct from dorsal part of vitelline reservoir to form short ovo-vitelline duct. Numerous fine gland ducts entering ovo-vitelline duct at base of proximal region of ootype (
<figureCitation id="1353B0D0D313FFE2FDB1408998B255D5" box="[601,673,1981,2007]" captionStart="FIGURE 6" captionStartId="12.[151,256,1736,1760]" captionTargetBox="[282,1310,205,1705]" captionTargetId="figure@12.[270,1326,194,1716]" captionTargetPageId="12" captionText="FIGURE 6. Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963. Anterolateral region of body (left side) of adult parasite. Composite drawing in ventral view based on AHC 29905 (specimen labeled no. 1) with some additions by hand. Note that positions of some ducts (e. g. transverse vitelline duct, vas deferens and uterus) are displaced slightly for clarity. Also, common genital papilla, common genital pore, excretory papilla and vaginal pore are located dorsally and are shown here as seen by transparency. Abbreviations: cgp, common genital papilla; cv, common vitelline duct; dv, distal part of vagina; ep, excretory papilla; ey, eye; fc, fertilisation chamber; ju, junction of uterus and terminal male genital duct; m, male accessory gland reservoir; mg, terminal male genital duct; og, ootype glands; ov, oviduct; ovo, ovovitelline duct; pv, proximal part of vagina; tvd, transverse vitelline duct; u, uterus; vd, vas deferens; vdc, vas deferens within cirrus sac; vr, vitelline reservoir. Other abbreviations as for Figure 1. Scale bar: 1 mm." httpUri="https://zenodo.org/record/196229/files/figure.png" pageId="11" pageNumber="12">Fig. 6</figureCitation>
); many small gland cells (not shown) located in curve of cirrus sac, right of oviduct. Ootype with convoluted proximal duct leading to tetrahedral-shaped chamber at level of excretory bladders, leading into uterus. Uterus long, following parallel course dorsal to terminal male genital duct, uniting near base of genital papilla (
<figureCitation id="1353B0D0D315FFE4FD6E42DF98DD5407" box="[646,718,1515,1541]" captionStart="FIGURE 6" captionStartId="12.[151,256,1736,1760]" captionTargetBox="[282,1310,205,1705]" captionTargetId="figure@12.[270,1326,194,1716]" captionTargetPageId="12" captionText="FIGURE 6. Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963. Anterolateral region of body (left side) of adult parasite. Composite drawing in ventral view based on AHC 29905 (specimen labeled no. 1) with some additions by hand. Note that positions of some ducts (e. g. transverse vitelline duct, vas deferens and uterus) are displaced slightly for clarity. Also, common genital papilla, common genital pore, excretory papilla and vaginal pore are located dorsally and are shown here as seen by transparency. Abbreviations: cgp, common genital papilla; cv, common vitelline duct; dv, distal part of vagina; ep, excretory papilla; ey, eye; fc, fertilisation chamber; ju, junction of uterus and terminal male genital duct; m, male accessory gland reservoir; mg, terminal male genital duct; og, ootype glands; ov, oviduct; ovo, ovovitelline duct; pv, proximal part of vagina; tvd, transverse vitelline duct; u, uterus; vd, vas deferens; vdc, vas deferens within cirrus sac; vr, vitelline reservoir. Other abbreviations as for Figure 1. Scale bar: 1 mm." httpUri="https://zenodo.org/record/196229/files/figure.png" pageId="13" pageNumber="14">Fig. 6</figureCitation>
; see also
<bibRefCitation id="EFF9D1A4D315FFE4FCA942DF9E955407" author="Kearn" box="[833,1158,1515,1541]" pageId="13" pageNumber="28" refString="Kearn, G. C. &amp; Whittington, I. D. (1992) Diversity of reproductive behaviour in platyhelminth parasites: insemination in some benedeniine (capsalid) monogeneans. Parasitology, 104, 489 - 496." type="journal article" year="1992">Kearn &amp; Whittington 1992</bibRefCitation>
). Common genital duct short, opening dorsomarginally at common genital pore at distal end of short genital papilla (
<figureCitation id="1353B0D0D315FFE4FB3E41249F335428" box="[1238,1312,1552,1578]" captionStart="FIGURE 1" captionStartId="6.[151,258,1755,1779]" captionTargetBox="[278,1303,216,1729]" captionTargetId="figure@6.[267,1321,197,1745]" captionTargetPageId="6" captionText="FIGURE 1. Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963. Whole adult parasite, composite drawing, in ventral view based on AHC 29906 (specimen labeled no. 1) with some additions by hand. For clarity, extent of intestinal diverticula (entirely coextensive with the vitellarium) other than immediately posterior to pharynx and separate uterus parallel with terminal male genital duct are not shown and extensive convolutions of vas deferens are simplified (but are shown in full in Figure 6). Abbreviations: aao, anterior attachment organ; ah, anterior hamulus; as, accessory sclerite; c, cirrus; cp, common genital pore; cs, cirrus sac; dh, dorsal horn; e, excretory bladder; em, extrinsic muscle; g, germarium; gr, ventral groove; h, haptor; ho, hooklet; i, intestine; mv, marginal valve; n, longitudinal nerve trunk; o, ootype; p, pharynx; ph, posterior hamulus; rp, rounded papilla; t, testis; te, tendon; v, vagina; vi, vitelline follicle; vp, vaginal pore. Scale bar: 1 mm." httpUri="https://zenodo.org/record/196224/files/figure.png" pageId="13" pageNumber="14">Figs 1</figureCitation>
,
<figureCitation id="1353B0D0D315FFE4FAC741249F2D5428" box="[1327,1342,1552,1578]" captionStart="FIGURE 4" captionStartId="9.[151,255,1360,1384]" captionTargetBox="[531,1055,180,1334]" captionTargetId="figure@9.[531,1056,176,1339]" captionTargetPageId="9" captionText="FIGURE 4. Scanning electron micrographs of dorsal region of juvenile specimen of Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963. A. Anterior region of specimen showing small tapering papillae resembling horns (h) dorsal to junction of body and anterior attachment organs. The position of the common genital pore (c) and the excretory papillae (ep) are also indicated. B. Haptor and posterior region of body proper showing pair of conical structures (co) near peduncle. Note marginal valve (mv). C. Enlargement of Figure 4 B showing pair of conical structures dorsal to haptor. Scale bars: A, 250 µm; B, 250 µm; C, 100 µm." httpUri="https://zenodo.org/record/196227/files/figure.png" pageId="13" pageNumber="14">4</figureCitation>
A, 6).
</paragraph>
<caption id="DF17FCDDD314FFE5FF7F41FC9E6A55DD" httpUri="https://zenodo.org/record/196229/files/figure.png" pageId="12" pageNumber="13" targetBox="[282,1310,205,1705]" targetPageId="12">
<paragraph id="8BD7AC55D314FFE5FF7F41FC9E6A55DD" blockId="12.[151,1437,1736,2015]" pageId="12" pageNumber="13">
<emphasis id="B91C7047D314FFE5FF7F41FC9B0F54E2" bold="true" box="[151,284,1736,1760]" pageId="12" pageNumber="13">FIGURE 6.</emphasis>
<taxonomicName id="4C68D7D6D314FFE5FECB41FD999A54DD" authority="(Luhe, 1906) Yamaguti, 1963" authorityName="(Luhe, 1906) Yamaguti" authorityYear="1963" box="[291,905,1736,1760]" class="Monogenea" family="Capsalidae" genus="Benedeniella" kingdom="Animalia" order="Capsalidea" pageId="12" pageNumber="13" phylum="Platyhelminthes" rank="species" species="macrocolpa">
<emphasis id="B91C7047D314FFE5FECB41FD982954E2" box="[291,570,1737,1760]" italics="true" pageId="12" pageNumber="13">Benedeniella macrocolpa</emphasis>
(Lühe, 1906) Yamaguti, 1963
</taxonomicName>
. Anterolateral region of body (left side) of adult parasite. Composite drawing in ventral view based on AHC 29905 (specimen labeled no. 1) with some additions by hand. Note that positions of some ducts (e.g. transverse vitelline duct, vas deferens and uterus) are displaced slightly for clarity. Also, common genital papilla, common genital pore, excretory papilla and vaginal pore are located dorsally and are shown here as seen by transparency. Abbreviations: cgp, common genital papilla; cv, common vitelline duct; dv, distal part of vagina; ep, excretory papilla; ey, eye; fc, fertilisation chamber; ju, junction of uterus and terminal male genital duct; m, male accessory gland reservoir; mg, terminal male genital duct; og, ootype glands; ov, oviduct; ovo, ovovitelline duct; pv, proximal part of vagina; tvd, transverse vitelline duct; u, uterus; vd, vas deferens; vdc, vas deferens within cirrus sac; vr, vitelline reservoir. Other abbreviations as for Figure 1. Scale bar: 1 mm.
</paragraph>
</caption>
<caption id="DF17FCDDD315FFE4FF7F43EF98975770" httpUri="https://zenodo.org/record/196230/files/figure.png" pageId="13" pageNumber="14" targetBox="[207,1398,209,1213]" targetPageId="13">
<paragraph id="8BD7AC55D315FFE4FF7F43EF98975770" blockId="13.[151,1436,1243,1394]" pageId="13" pageNumber="14">
<emphasis id="B91C7047D315FFE4FF7F43EF9B0F56F1" bold="true" box="[151,284,1243,1267]" pageId="13" pageNumber="14">FIGURE 7.</emphasis>
Diagrammatic comparison of curvature of cirrus sac in example juvenile (A) and adult (B)
<taxonomicName id="4C68D7D6D315FFE4FAE643EF987A5710" authority="(Luhe, 1906) Yamaguti, 1963" authorityName="(Luhe, 1906) Yamaguti" authorityYear="1963" class="Monogenea" family="Capsalidae" genus="Benedeniella" kingdom="Animalia" order="Capsalidea" pageId="13" pageNumber="14" phylum="Platyhelminthes" rank="species" species="macrocolpa">
<emphasis id="B91C7047D315FFE4FAE643EF9B0A5710" italics="true" pageId="13" pageNumber="14">Benedeniella macrocolpa</emphasis>
(Lühe, 1906) Yamaguti, 1963
</taxonomicName>
.
<emphasis id="B91C7047D315FFE4FD9D43CF989C5711" bold="true" box="[629,655,1275,1299]" pageId="13" pageNumber="14">A.</emphasis>
Juvenile specimen (AHC 29905 (specimen labeled no. 9); total length (including haptor) = 3.37 mm; anterior hamulus lengths = 366 and 376 µm.
<emphasis id="B91C7047D315FFE4FC23422F99F75731" bold="true" box="[971,996,1307,1331]" pageId="13" pageNumber="14">B.</emphasis>
Adult specimen (AHC 29905 (specimen labeled no. 8)); total length (including haptor) = 8.1 mm; anterior hamulus lengths = 718 and 743 µm. Abbreviations as for Figure 1. Scale bars: A, 300 µm; B, 1 mm.
</paragraph>
</caption>
<paragraph id="8BD7AC55D315FFE4FF2D41029F115578" blockId="13.[151,1437,1440,2026]" pageId="13" pageNumber="14">
Vitelline reservoir ventral, curving anteromedianly, near proximal end of cirrus sac. Vagina exceedingly long, 2.1–11 (6.6, n = 13, measured along curves) mm; originating proximally via narrow duct from left dorsal margin of vitelline reservoir. No seminal receptacles or proximal specialisations. Vagina following path posteriorly to a level well behind testes before turning anteriorly through 180°, passing ventral to left excretory bladder, then running parallel, just posterior to terminal male genital duct and uterus, to open dorsomarginally, just posterior to common genital papilla (
<figureCitation id="1353B0D0D315FFE4FCBE41C499B15508" box="[854,930,1776,1802]" captionStart="FIGURE 1" captionStartId="6.[151,258,1755,1779]" captionTargetBox="[278,1303,216,1729]" captionTargetId="figure@6.[267,1321,197,1745]" captionTargetPageId="6" captionText="FIGURE 1. Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963. Whole adult parasite, composite drawing, in ventral view based on AHC 29906 (specimen labeled no. 1) with some additions by hand. For clarity, extent of intestinal diverticula (entirely coextensive with the vitellarium) other than immediately posterior to pharynx and separate uterus parallel with terminal male genital duct are not shown and extensive convolutions of vas deferens are simplified (but are shown in full in Figure 6). Abbreviations: aao, anterior attachment organ; ah, anterior hamulus; as, accessory sclerite; c, cirrus; cp, common genital pore; cs, cirrus sac; dh, dorsal horn; e, excretory bladder; em, extrinsic muscle; g, germarium; gr, ventral groove; h, haptor; ho, hooklet; i, intestine; mv, marginal valve; n, longitudinal nerve trunk; o, ootype; p, pharynx; ph, posterior hamulus; rp, rounded papilla; t, testis; te, tendon; v, vagina; vi, vitelline follicle; vp, vaginal pore. Scale bar: 1 mm." httpUri="https://zenodo.org/record/196224/files/figure.png" pageId="13" pageNumber="14">Figs 1</figureCitation>
,
<figureCitation id="1353B0D0D315FFE4FC5941C499D05508" box="[945,963,1776,1802]" captionStart="FIGURE 6" captionStartId="12.[151,256,1736,1760]" captionTargetBox="[282,1310,205,1705]" captionTargetId="figure@12.[270,1326,194,1716]" captionTargetPageId="12" captionText="FIGURE 6. Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963. Anterolateral region of body (left side) of adult parasite. Composite drawing in ventral view based on AHC 29905 (specimen labeled no. 1) with some additions by hand. Note that positions of some ducts (e. g. transverse vitelline duct, vas deferens and uterus) are displaced slightly for clarity. Also, common genital papilla, common genital pore, excretory papilla and vaginal pore are located dorsally and are shown here as seen by transparency. Abbreviations: cgp, common genital papilla; cv, common vitelline duct; dv, distal part of vagina; ep, excretory papilla; ey, eye; fc, fertilisation chamber; ju, junction of uterus and terminal male genital duct; m, male accessory gland reservoir; mg, terminal male genital duct; og, ootype glands; ov, oviduct; ovo, ovovitelline duct; pv, proximal part of vagina; tvd, transverse vitelline duct; u, uterus; vd, vas deferens; vdc, vas deferens within cirrus sac; vr, vitelline reservoir. Other abbreviations as for Figure 1. Scale bar: 1 mm." httpUri="https://zenodo.org/record/196229/files/figure.png" pageId="13" pageNumber="14">6</figureCitation>
). Vitellarium follicular, extending from between anterior attachment organs almost to posterior extremity of body proper, coextensive with intestinal caeca (
<figureCitation id="1353B0D0D315FFE4FF0F400F9B3F5557" box="[231,300,1851,1877]" captionStart="FIGURE 1" captionStartId="6.[151,258,1755,1779]" captionTargetBox="[278,1303,216,1729]" captionTargetId="figure@6.[267,1321,197,1745]" captionTargetPageId="6" captionText="FIGURE 1. Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963. Whole adult parasite, composite drawing, in ventral view based on AHC 29906 (specimen labeled no. 1) with some additions by hand. For clarity, extent of intestinal diverticula (entirely coextensive with the vitellarium) other than immediately posterior to pharynx and separate uterus parallel with terminal male genital duct are not shown and extensive convolutions of vas deferens are simplified (but are shown in full in Figure 6). Abbreviations: aao, anterior attachment organ; ah, anterior hamulus; as, accessory sclerite; c, cirrus; cp, common genital pore; cs, cirrus sac; dh, dorsal horn; e, excretory bladder; em, extrinsic muscle; g, germarium; gr, ventral groove; h, haptor; ho, hooklet; i, intestine; mv, marginal valve; n, longitudinal nerve trunk; o, ootype; p, pharynx; ph, posterior hamulus; rp, rounded papilla; t, testis; te, tendon; v, vagina; vi, vitelline follicle; vp, vaginal pore. Scale bar: 1 mm." httpUri="https://zenodo.org/record/196224/files/figure.png" pageId="13" pageNumber="14">Fig. 1</figureCitation>
). Free eggs not observed; in some mounted specimens, eggs observed in ootype, with tetrahedral shape and appendage; no egg measurements made due to distortion during specimen processing.
</paragraph>
</subSubSection>
<subSubSection id="C372FFDED315FFE7FF2E40B19F9B5655" lastPageId="14" lastPageNumber="15" pageId="13" pageNumber="14" type="discussion">
<paragraph id="8BD7AC55D315FFE7FF2E40B199785320" blockId="13.[151,1437,1440,2026]" lastBlockId="14.[151,1437,152,1111]" lastPageId="14" lastPageNumber="15" pageId="13" pageNumber="14">
<emphasis id="B91C7047D315FFE4FF2E40B19B51559D" bold="true" box="[198,322,1925,1951]" pageId="13" pageNumber="14">Remarks:</emphasis>
Live specimens of
<taxonomicName id="4C68D7D6D315FFE4FDC340B298C4559D" box="[555,727,1926,1951]" class="Monogenea" family="Capsalidae" genus="Benedeniella" kingdom="Animalia" order="Capsalidea" pageId="13" pageNumber="14" phylum="Platyhelminthes" rank="species" species="macrocolpa">
<emphasis id="B91C7047D315FFE4FDC340B298C4559D" box="[555,727,1926,1951]" italics="true" pageId="13" pageNumber="14">B. macrocolpa</emphasis>
</taxonomicName>
from
<taxonomicName id="4C68D7D6D315FFE4FCC840B299B6559D" box="[800,933,1926,1951]" class="Elasmobranchii" family="Myliobatidae" genus="Rhinoptera" kingdom="Animalia" order="Myliobatiformes" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="neglecta">
<emphasis id="B91C7047D315FFE4FCC840B299B6559D" box="[800,933,1926,1951]" italics="true" pageId="13" pageNumber="14">R. neglecta</emphasis>
</taxonomicName>
collected from Moreton Bay, Queensland,
<collectingCountry id="F37FECC5D315FFE4FF7F409F9B1855C7" box="[151,267,1963,1989]" name="Australia" pageId="13" pageNumber="14">Australia</collectingCountry>
were apparently healthy. They attached to the bottom of glass Petri dishes containing FSW, displayed leech-like locomotion by alternately attaching the haptor and anterior attachment organs (e.g. see
<bibRefCitation id="EFF9D1A4D316FFE7FF7F47AC9BA352B0" author="Whittington" box="[151,432,152,178]" pageId="14" pageNumber="29" refString="Whittington, I. D., Cribb, B. W., Hamwood, T. E. &amp; Halliday, J. A. (2000) Host-specificity of monogenean (platyhelminth) parasites: a role for anterior adhesive areas? International Journal for Parasitology, 30, 305 - 320." type="journal article" year="2000">
Whittington
<emphasis id="B91C7047D316FFE7FEC647AC9B7052B3" box="[302,355,152,177]" italics="true" pageId="14" pageNumber="15">et al</emphasis>
. 2000
</bibRefCitation>
) and also exhibited characteristic undulating movements of the body while attached by the haptor. For the capsalid
<taxonomicName id="4C68D7D6D316FFE7FDC1478999DF52D5" authority="Kearn (1962)" authorityName="Kearn" authorityYear="1962" box="[553,972,189,215]" class="Monogenea" family="Capsalidae" genus="Entobdella" kingdom="Animalia" order="Capsalidea" pageId="14" pageNumber="15" phylum="Platyhelminthes" rank="species" species="soleae">
<emphasis id="B91C7047D316FFE7FDC14789990352D4" box="[553,784,189,214]" italics="true" pageId="14" pageNumber="15">Entobdella soleae</emphasis>
,
<bibRefCitation id="EFF9D1A4D316FFE7FCC9478999DF52D5" author="Kearn" box="[801,972,189,215]" pageId="14" pageNumber="28" refString="Kearn, G. C. (1962) Breathing movements in Entobdella soleae (Trematoda, Monogenea) from the skin of the common sole. Journal of the Marine Biological Association of the United Kingdom, 42, 93 - 104." type="journal article" year="1962">Kearn (1962)</bibRefCitation>
</taxonomicName>
has demonstrated that these regular undulatory motions probably aid respiration of the parasite. Living specimens of
<taxonomicName id="4C68D7D6D316FFE7FB8B47D69F0252F9" box="[1123,1297,226,251]" class="Monogenea" family="Capsalidae" genus="Benedeniella" kingdom="Animalia" order="Capsalidea" pageId="14" pageNumber="15" phylum="Platyhelminthes" rank="species" species="macrocolpa">
<emphasis id="B91C7047D316FFE7FB8B47D69F0252F9" box="[1123,1297,226,251]" italics="true" pageId="14" pageNumber="15">B. macrocolpa</emphasis>
</taxonomicName>
attached to glass dishes failed to lay any eggs during periods of up to 7 h.
</paragraph>
<paragraph id="8BD7AC55D316FFE7FF2E461998905025" blockId="14.[151,1437,152,1111]" pageId="14" pageNumber="15">
The anatomy of
<taxonomicName id="4C68D7D6D316FFE7FE6D461998225344" box="[389,561,301,326]" class="Monogenea" family="Capsalidae" genus="Benedeniella" kingdom="Animalia" order="Capsalidea" pageId="14" pageNumber="15" phylum="Platyhelminthes" rank="species" species="macrocolpa">
<emphasis id="B91C7047D316FFE7FE6D461998225344" box="[389,561,301,326]" italics="true" pageId="14" pageNumber="15">B. macrocolpa</emphasis>
</taxonomicName>
redescribed here is consistent in most respects with that presented by
<bibRefCitation id="EFF9D1A4D316FFE7FA8846199AFA536E" author="Luhe" pageId="14" pageNumber="29" refString="Luhe, M. (1906) Report on the trematode parasites from the marine fishes of Ceylon. Report to the Government of Ceylon on the Pearl Oyster Fisheries of the Gulf of Manaar, Supplementary Report, 5, 97 - 108." type="journal article" year="1906">Lühe (1906)</bibRefCitation>
. The only major difference relates to the arrangement of the terminal female and male genital systems.
<bibRefCitation id="EFF9D1A4D316FFE7FF7F464C9B205390" author="Luhe" box="[151,307,376,402]" pageId="14" pageNumber="29" refString="Luhe, M. (1906) Report on the trematode parasites from the marine fishes of Ceylon. Report to the Government of Ceylon on the Pearl Oyster Fisheries of the Gulf of Manaar, Supplementary Report, 5, 97 - 108." type="journal article" year="1906">Lühe (1906)</bibRefCitation>
described a short uterus that opened into a long common genital duct. The present study, however, confirms the view of
<bibRefCitation id="EFF9D1A4D316FFE7FDE146A9994F53B5" author="Kearn" box="[521,860,413,439]" pageId="14" pageNumber="28" refString="Kearn, G. C. &amp; Whittington, I. D. (1992) Diversity of reproductive behaviour in platyhelminth parasites: insemination in some benedeniine (capsalid) monogeneans. Parasitology, 104, 489 - 496." type="journal article" year="1992">Kearn &amp; Whittington (1992)</bibRefCitation>
that the uterus of
<taxonomicName id="4C68D7D6D316FFE7FBDD46A99EF153B4" box="[1077,1250,413,438]" class="Monogenea" family="Capsalidae" genus="Benedeniella" kingdom="Animalia" order="Capsalidea" pageId="14" pageNumber="15" phylum="Platyhelminthes" rank="species" species="macrocolpa">
<emphasis id="B91C7047D316FFE7FBDD46A99EF153B4" box="[1077,1250,413,438]" italics="true" pageId="14" pageNumber="15">B. macrocolpa</emphasis>
</taxonomicName>
is, in fact, long and runs parallel with and dorsal to the separate elongate terminal male genital duct. The female and male systems unite only distally near the base of a short common genital papilla to form a short common genital duct which opens dorsally (
<figureCitation id="1353B0D0D316FFE7FE31453998305025" box="[473,547,525,551]" captionStart="FIGURE 1" captionStartId="6.[151,258,1755,1779]" captionTargetBox="[278,1303,216,1729]" captionTargetId="figure@6.[267,1321,197,1745]" captionTargetPageId="6" captionText="FIGURE 1. Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963. Whole adult parasite, composite drawing, in ventral view based on AHC 29906 (specimen labeled no. 1) with some additions by hand. For clarity, extent of intestinal diverticula (entirely coextensive with the vitellarium) other than immediately posterior to pharynx and separate uterus parallel with terminal male genital duct are not shown and extensive convolutions of vas deferens are simplified (but are shown in full in Figure 6). Abbreviations: aao, anterior attachment organ; ah, anterior hamulus; as, accessory sclerite; c, cirrus; cp, common genital pore; cs, cirrus sac; dh, dorsal horn; e, excretory bladder; em, extrinsic muscle; g, germarium; gr, ventral groove; h, haptor; ho, hooklet; i, intestine; mv, marginal valve; n, longitudinal nerve trunk; o, ootype; p, pharynx; ph, posterior hamulus; rp, rounded papilla; t, testis; te, tendon; v, vagina; vi, vitelline follicle; vp, vaginal pore. Scale bar: 1 mm." httpUri="https://zenodo.org/record/196224/files/figure.png" pageId="14" pageNumber="15">Figs 1</figureCitation>
,
<figureCitation id="1353B0D0D316FFE7FDD94539982C5025" box="[561,575,525,551]" captionStart="FIGURE 5" captionStartId="10.[151,259,1317,1341]" captionTargetBox="[250,1335,504,1289]" captionTargetId="figure@10.[243,1344,493,1297]" captionTargetPageId="10" captionText="FIGURE 5. Diagrammatic outlines of adult Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963 (A) and B. posterocolpa (Hargis, 1955) Yamaguti, 1963 (B) in dorsal view showing relative positions of paired anterior tapering horns (dh), common genital papilla (cgp; absent in B. posterocolpa), paired excretory papillae (ep) and paired conical structures (co). Drawn from several wholemounted specimens viewed dorsally. Scale bar: 4 mm." httpUri="https://zenodo.org/record/196228/files/figure.png" pageId="14" pageNumber="15">5</figureCitation>
A, 6).
</paragraph>
<paragraph id="8BD7AC55D316FFE7FF2E45069EF15175" blockId="14.[151,1437,152,1111]" pageId="14" pageNumber="15">
Additional anatomical features emphasised in this redescription of
<taxonomicName id="4C68D7D6D316FFE7FC0F45069E865049" box="[999,1173,562,587]" class="Monogenea" family="Capsalidae" genus="Benedeniella" kingdom="Animalia" order="Capsalidea" pageId="14" pageNumber="15" phylum="Platyhelminthes" rank="species" species="macrocolpa">
<emphasis id="B91C7047D316FFE7FC0F45069E865049" box="[999,1173,562,587]" italics="true" pageId="14" pageNumber="15">B. macrocolpa</emphasis>
</taxonomicName>
relate to the presence and path of conspicuous tendons from extrinsic muscles in the body associated with the haptor and a series of 11–28 complete roughly parallel, ventral, narrow grooves on the anterolateral margin of each anterior attachment organ. In some specimens, there may be some incomplete grooves between the complete grooves. The grooves on the anterior attachment organs of
<taxonomicName id="4C68D7D6D316FFE7FD1045FC99BA50E3" box="[760,937,712,737]" class="Monogenea" family="Capsalidae" genus="Benedeniella" kingdom="Animalia" order="Capsalidea" pageId="14" pageNumber="15" phylum="Platyhelminthes" rank="species" species="macrocolpa">
<emphasis id="B91C7047D316FFE7FD1045FC99BA50E3" box="[760,937,712,737]" italics="true" pageId="14" pageNumber="15">B. macrocolpa</emphasis>
</taxonomicName>
resemble similar structures described by
<bibRefCitation id="EFF9D1A4D316FFE7FF7F45D99B535105" author="Hargis" box="[151,320,749,775]" pageId="14" pageNumber="28" refString="Hargis, W. J. Jr. (1955) Monogenetic trematodes of Gulf of Mexico fishes. Part V. The superfamily Capsaloidea. Transactions of the American Microscopical Society, 74, 203 - 225." type="journal article" year="1955">Hargis (1955)</bibRefCitation>
on the anterior attachment organs of
<taxonomicName id="4C68D7D6D316FFE7FCE145D999DA5104" box="[777,969,749,774]" class="Monogenea" family="Capsalidae" genus="Benedenia" kingdom="Animalia" order="Capsalidea" pageId="14" pageNumber="15" phylum="Platyhelminthes" rank="species" species="posterocolpa">
<emphasis id="B91C7047D316FFE7FCE145D999DA5104" box="[777,969,749,774]" italics="true" pageId="14" pageNumber="15">B. posterocolpa</emphasis>
</taxonomicName>
and their presence is confirmed in the redescription below. The route of the tendons in the haptor of
<taxonomicName id="4C68D7D6D316FFE7FC9444269E3A5129" box="[892,1065,786,811]" class="Monogenea" family="Capsalidae" genus="Benedeniella" kingdom="Animalia" order="Capsalidea" pageId="14" pageNumber="15" phylum="Platyhelminthes" rank="species" species="macrocolpa">
<emphasis id="B91C7047D316FFE7FC9444269E3A5129" box="[892,1065,786,811]" italics="true" pageId="14" pageNumber="15">B. macrocolpa</emphasis>
</taxonomicName>
and their attachment mostly to the proximal ends of the anterior hamuli is shared by
<taxonomicName id="4C68D7D6D316FFE7FCE5440C99D65153" box="[781,965,824,849]" class="Monogenea" family="Capsalidae" genus="Benedenia" kingdom="Animalia" order="Capsalidea" pageId="14" pageNumber="15" phylum="Platyhelminthes" rank="species" species="posterocolpa">
<emphasis id="B91C7047D316FFE7FCE5440C99D65153" box="[781,965,824,849]" italics="true" pageId="14" pageNumber="15">B. posterocolpa</emphasis>
</taxonomicName>
(see below),
<taxonomicName id="4C68D7D6D316FFE7FB8D440C9EF55153" box="[1125,1254,824,849]" class="Monogenea" family="Capsalidae" genus="Entobdella" kingdom="Animalia" order="Capsalidea" pageId="14" pageNumber="15" phylum="Platyhelminthes" rank="genus">
<emphasis id="B91C7047D316FFE7FB8D440C9EF55153" box="[1125,1254,824,849]" italics="true" pageId="14" pageNumber="15">Entobdella</emphasis>
</taxonomicName>
species (e.g. in
<taxonomicName id="4C68D7D6D316FFE7FF7F44699B115174" box="[151,258,861,886]" class="Monogenea" family="Capsalidae" genus="Entobdella" kingdom="Animalia" order="Capsalidea" pageId="14" pageNumber="15" phylum="Platyhelminthes" rank="species" species="soleae">
<emphasis id="B91C7047D316FFE7FF7F44699B115174" box="[151,258,861,886]" italics="true" pageId="14" pageNumber="15">E. soleae</emphasis>
</taxonomicName>
, see
<bibRefCitation id="EFF9D1A4D316FFE7FED544699BDA5175" author="Kearn" box="[317,457,861,887]" pageId="14" pageNumber="28" refString="Kearn, G. C. (1964) The attachment of the monogenean Entobdella soleae to the skin of the common sole. Parasitology, 54, 327 - 335." type="journal article" year="1964">Kearn 1964</bibRefCitation>
) and some
<taxonomicName id="4C68D7D6D316FFE7FDA7446998E95174" box="[591,762,861,886]" class="Monogenea" family="Capsalidae" genus="Neoentobdella" kingdom="Animalia" order="Capsalidea" pageId="14" pageNumber="15" phylum="Platyhelminthes" rank="genus">
<emphasis id="B91C7047D316FFE7FDA7446998E95174" box="[591,762,861,886]" italics="true" pageId="14" pageNumber="15">Neoentobdella</emphasis>
</taxonomicName>
species (see
<bibRefCitation id="EFF9D1A4D316FFE7FC7B44699EC65175" author="Kearn" box="[915,1237,861,887]" pageId="14" pageNumber="28" refString="Kearn, G. C. &amp; Whittington, I. D. (2005) Neoentobdella gen. nov. for species of Entobdella Blainville in Lamarck, 1818 (Monogenea, Capsalidae, Entobdellinae) from stingray hosts, with descriptions of two new species. Acta Parasitologica, 50, 32 - 48." type="journal article" year="2005">Kearn &amp; Whittington 2005</bibRefCitation>
).
</paragraph>
<paragraph id="8BD7AC55D316FFE7FF2E44B69F9B5655" blockId="14.[151,1437,152,1111]" pageId="14" pageNumber="15">
This redescription of
<taxonomicName id="4C68D7D6D316FFE7FE3144B6989F5199" box="[473,652,898,923]" class="Monogenea" family="Capsalidae" genus="Benedeniella" kingdom="Animalia" order="Capsalidea" pageId="14" pageNumber="15" phylum="Platyhelminthes" rank="species" species="macrocolpa">
<emphasis id="B91C7047D316FFE7FE3144B6989F5199" box="[473,652,898,923]" italics="true" pageId="14" pageNumber="15">B. macrocolpa</emphasis>
</taxonomicName>
has also identified dorsal structures associated, in anterior to posterior succession, with the anterior attachment organs, common genital pore, excretory bladders and at the junction of the body with the haptor. The anteriormost tapering papillae are horn-like and were also described by
<bibRefCitation id="EFF9D1A4D316FFE7FF5444C69B9C560E" author="McMahon" box="[188,399,1010,1036]" pageId="14" pageNumber="29" refString="McMahon, J. W. (1963) Monogenetic trematodes from some Chesapeake Bay fishes. Part I: The superfamilies Capsaloidea Price, 1936 and Diclidophoroidea Price, 1936. Chesapeake Science, 4, 151 - 160." type="journal article" year="1963">McMahon (1963)</bibRefCitation>
for
<taxonomicName id="4C68D7D6D316FFE7FE2A44C6986E5609" box="[450,637,1010,1035]" class="Monogenea" family="Capsalidae" genus="Benedenia" kingdom="Animalia" order="Capsalidea" pageId="14" pageNumber="15" phylum="Platyhelminthes" rank="species" species="posterocolpa">
<emphasis id="B91C7047D316FFE7FE2A44C6986E5609" box="[450,637,1010,1035]" italics="true" pageId="14" pageNumber="15">B. posterocolpa</emphasis>
</taxonomicName>
(see below). With the exception of the common genital papilla, all dorsal structures identified here for
<taxonomicName id="4C68D7D6D316FFE7FDB0432C991D5633" box="[600,782,1048,1073]" class="Monogenea" family="Capsalidae" genus="Benedeniella" kingdom="Animalia" order="Capsalidea" pageId="14" pageNumber="15" phylum="Platyhelminthes" rank="species" species="macrocolpa">
<emphasis id="B91C7047D316FFE7FDB0432C991D5633" box="[600,782,1048,1073]" italics="true" pageId="14" pageNumber="15">B. macrocolpa</emphasis>
</taxonomicName>
have also been identified in the present study for
<taxonomicName id="4C68D7D6D316FFE7FA6A432C9B3C5654" class="Monogenea" family="Capsalidae" genus="Benedenia" kingdom="Animalia" order="Capsalidea" pageId="14" pageNumber="15" phylum="Platyhelminthes" rank="species" species="posterocolpa">
<emphasis id="B91C7047D316FFE7FA6A432C9B3C5654" italics="true" pageId="14" pageNumber="15">B. posterocolpa</emphasis>
</taxonomicName>
(see
<figureCitation id="1353B0D0D316FFE7FE8443099BA25655" box="[364,433,1085,1111]" captionStart="FIGURE 5" captionStartId="10.[151,259,1317,1341]" captionTargetBox="[250,1335,504,1289]" captionTargetId="figure@10.[243,1344,493,1297]" captionTargetPageId="10" captionText="FIGURE 5. Diagrammatic outlines of adult Benedeniella macrocolpa (Lühe, 1906) Yamaguti, 1963 (A) and B. posterocolpa (Hargis, 1955) Yamaguti, 1963 (B) in dorsal view showing relative positions of paired anterior tapering horns (dh), common genital papilla (cgp; absent in B. posterocolpa), paired excretory papillae (ep) and paired conical structures (co). Drawn from several wholemounted specimens viewed dorsally. Scale bar: 4 mm." httpUri="https://zenodo.org/record/196228/files/figure.png" pageId="14" pageNumber="15">Fig. 5</figureCitation>
and below). These characters are included in the amended generic diagnosis above.
</paragraph>
</subSubSection>
</treatment>
</document>
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