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<mods:title id="1162D06A7E81D56C2156F3DD49D611A6">A revision of Entobdella Blainville in Lamarck, 1818, with special emphasis on the nominal (type) species “ Entobdella hippoglossi (Müller, 1776) Blainville, 1818 ” (Monogenea: Capsalidae: Entobdellinae) from teleost flatfishes, with descriptions of three new species and a new genus</mods:title>
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<mods:affiliation id="388BD56534EA127343BC1F9A1E304927">School of Biological Sciences, University of East Anglia, Norwich, NR 4 7 TJ, Norfolk, U. K. E-mail: g. kearn @ uea. ac. uk; r. evansgowing @ uea. ac. uk Monogenean Research Laboratory, Parasitology Section, The South Australian Museum, North Terrace, Adelaide, South Australia 5000, Australia. E-mail: whittington. ian @ saugov. sa. gov. au Marine Parasitology Laboratory, School of Earth and Environmental Sciences (DX 650 418), The University of Adelaide, North Terrace, Adelaide, South Australia 5005, Australia</mods:affiliation>
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<mods:namePart id="D3076DF779D77FEBB4F1D75C127E2D83">Evans-Gowing, Richard</mods:namePart>
<mods:affiliation id="539F75097A4690036E0A4FBE8BA47C46">School of Biological Sciences, University of East Anglia, Norwich, NR 4 7 TJ, Norfolk, U. K. E-mail: g. kearn @ uea. ac. uk; r. evansgowing @ uea. ac. uk Monogenean Research Laboratory, Parasitology Section, The South Australian Museum, North Terrace, Adelaide, South Australia 5000, Australia. E-mail: whittington. ian @ saugov. sa. gov. au Marine Parasitology Laboratory, School of Earth and Environmental Sciences (DX 650 418), The University of Adelaide, North Terrace, Adelaide, South Australia 5005, Australia</mods:affiliation>
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<paragraph id="D04536E6FF94FF83159FF8F9FC7CB15B" blockId="21.[151,934,1815,1876]" box="[151,934,1815,1841]" pageId="21" pageNumber="22">
<heading id="8B0D818AFF94FF83159FF8F9FC7CB15B" bold="true" box="[151,934,1815,1841]" fontSize="11" level="1" pageId="21" pageNumber="22" reason="1">
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<taxonomicName id="17FA4D65FF94FF83159FF8F9FEB7B15B" ID-CoL="39YP6" baseAuthorityName="Van Beneden &amp; Hesse" baseAuthorityYear="1863" box="[151,365,1815,1841]" class="Monogenea" family="Capsalidae" genus="Entobdella" kingdom="Animalia" order="Capsalidea" pageId="21" pageNumber="22" phylum="Platyhelminthes" rank="species" species="soleae">
<emphasis id="E28EEAF4FF94FF83159FF8F9FEB7B15B" bold="true" box="[151,365,1815,1841]" italics="true" pageId="21" pageNumber="22">Entobdella soleae</emphasis>
</taxonomicName>
(
<bibRefCitation id="B46B4B17FF94FF831474F8F9FD01B15B" author="Van Beneden, P. - J. &amp; Hesse, E." box="[380,731,1815,1841]" pageId="21" pageNumber="22" pagination="1 - 142" refId="ref37134" refString="Van Beneden, P. - J. &amp; Hesse, E. (1864) Recherches sur les bdellodes (hirudinees) et les trematodes marins. Memoires de l'Academie Royale des Sciences, des Lettres et des Beaux-arts de Belgique 34, 1 - 142." type="journal article" year="1864">
Van Beneden &amp; 
<collectingRegion id="123EF804FF94FF831741F8F9FD48B15B" box="[585,658,1815,1841]" country="Germany" name="Hessen" pageId="21" pageNumber="22">Hesse</collectingRegion>
, 1864
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) Johnston, 1929
</emphasis>
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</paragraph>
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<subSubSection id="98E0656DFF94FF80159FF8D4FD9FB1B2" lastPageId="22" lastPageNumber="23" pageId="21" pageNumber="22" type="description">
<paragraph id="D04536E6FF94FF83159FF8D4FE42B13E" blockId="21.[151,934,1815,1876]" box="[151,408,1850,1876]" pageId="21" pageNumber="22">
(
<figureCitation id="48C12A63FF94FF831597F8D4FED6B13E" box="[159,268,1850,1876]" captionStart="FIGURE 15" captionStartId="15.[151,255,1634,1658]" captionTargetBox="[215,1390,834,1594]" captionTargetId="figure-365@15.[184,1411,794,1641]" captionTargetPageId="15" captionText="FIGURE 15. Entobdella spp. Accessory sclerites of adults. A. “E. hippoglossi type 1” from H. hippoglossus. B. “E. hippoglossi type 2” from H. hippoglossus. C. “Entobdella hippoglossi” from H. stenolepis. D. E. steingroeveri. E. E. squamula from Paralichthys californicus. F, G. accessory sclerites from 2 individuals of different sizes (ages) of a proposed new species from Hippoglossina macrops (F, Entobdella sp. 1 from Dr. M.T. González; G, from LUCE; see Table 2). H. E. brattstroemi from P. adspersus (Entobdella sp. 2 from Dr M.T. González; Table 2). J. E. soleae from Solea solea. K. E. rosaceus from Zalembius rosaceus. All sclerites drawn to the same scale with the distal region uppermost. Scale bars: 100µm." pageId="21" pageNumber="22">Figs. 15J</figureCitation>
, 
<figureCitation id="48C12A63FF94FF831411F8D4FEBEB13E" box="[281,356,1850,1876]" captionStart="FIGURES 27 – 31" captionStartId="24.[151,269,1644,1668]" captionTargetBox="[264,1323,211,1585]" captionTargetId="figure-160@24.[264,1323,194,1585]" captionTargetPageId="24" captionText="FIGURES 27 – 31. Entobdella soleae. Light microscope photographs of toluidine-blue-stained serial resin sections through the vaginal region of an adult specimen. 27. Extended section showing the ventrally situated vaginal opening (arrow) in relation to the germarium (g). 28. Section through the narrow, distal entrance tube (et) of the vagina. 29. Section through the narrow, distal entrance tube (et) and 1 of the coils of the expanded proximal region (pr) of the vagina. 30. Section through the junction between the narrow entrance tube (et) and the expanded first coil of the proximal region (pr) of the vagina. 31. Extended section showing spacious coils of the proximal region (pr) of the vagina in relation to the vitelline reservoir (vr) and the germarium (g). Heavily stained vertical lines at right angles to the dorsal surface are folds in the resin section. Scale bars: 50µm." pageId="21" pageNumber="22">27–31</figureCitation>
, 
<figureCitation id="48C12A63FF94FF831479F8D4FE55B13E" box="[369,399,1850,1876]" captionStart="FIGURE 37" captionStartId="32.[151,255,1897,1921]" captionTargetBox="[226,1392,197,1863]" captionTargetId="figure-76@32.[180,1424,146,1896]" captionTargetPageId="32" captionText="FIGURE 37. Entobdella soleae. A. Whole parasite in ventral view showing haptor papillae. Distribution of papillae derived from an SEM photograph by Dr J.A. Rees (see Kearn, 1998, fig. 4.3). B. Accessory sclerite enlarged. Abbreviation: cf, circular fibres associated with vaginal opening. Other abbreviations as in Figs. 26, 32. Scale bars: A, 1mm; B, 100µm." pageId="21" pageNumber="22">37</figureCitation>
)
</paragraph>
<paragraph id="D04536E6FF94FF83159FF864FBEBB19E" blockId="21.[151,1436,1930,2036]" pageId="21" pageNumber="22">
One of us (GCK) has studied the anatomy, ultrastructure and biology of 
<taxonomicName id="17FA4D65FF94FF8316FDF865FBB8B1CE" baseAuthorityName="Van Beneden &amp; Hesse" baseAuthorityYear="1863" box="[1013,1122,1931,1956]" class="Monogenea" family="Capsalidae" genus="Entobdella" kingdom="Animalia" order="Capsalidea" pageId="21" pageNumber="22" phylum="Platyhelminthes" rank="species" species="soleae">
<emphasis id="E28EEAF4FF94FF8316FDF865FBB8B1CE" box="[1013,1122,1931,1956]" italics="true" pageId="21" pageNumber="22">E. soleae</emphasis>
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from the skin of the common sole 
<taxonomicName id="17FA4D65FF94FF831401F85DFE55B1A6" authority="solea" authorityName="solea (Linnaeus" authorityYear="1758" box="[265,399,1971,1996]" class="Actinopterygii" family="Soleidae" genus="Solea" kingdom="Animalia" order="Pleuronectiformes" pageId="21" pageNumber="22" phylum="Chordata" rank="species" species="solea">
<emphasis id="E28EEAF4FF94FF831401F85DFE55B1A6" box="[265,399,1971,1996]" italics="true" pageId="21" pageNumber="22">Solea solea</emphasis>
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extensively (see Kearn 1998, 2004). Some points of relevance to the current project and some new observations on sections of resin-embedded material are given here.
</paragraph>
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<paragraph id="D04536E6FF97FF80159FFC53FE3FB59D" blockId="22.[151,1436,957,1015]" pageId="22" pageNumber="23">
<emphasis id="E28EEAF4FF97FF80159FFC53FEFCB5BF" bold="true" box="[151,294,957,981]" pageId="22" pageNumber="23">FIGURE 25.</emphasis>
<taxonomicName id="17FA4D65FF97FF801424FC50FDEEB5BF" authorityName="Johnston" authorityYear="1929" baseAuthorityName="Cohn" baseAuthorityYear="1916" box="[300,564,958,981]" class="Monogenea" family="Capsalidae" genus="Entobdella" kingdom="Animalia" order="Capsalidea" pageId="22" pageNumber="23" phylum="Platyhelminthes" rank="species" species="steingroeveri">
<emphasis id="E28EEAF4FF97FF801424FC50FDEEB5BF" box="[300,564,958,981]" italics="true" pageId="22" pageNumber="23">Entobdella steingroeveri</emphasis>
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. SEM photograph of the anterior region of the ventral surface of the haptor showing papillae. Scale bar: 200µm.
</paragraph>
</caption>
<paragraph id="D04536E6FF97FF8015CFFBC8FD14B2FA" blockId="22.[151,1436,1062,2008]" pageId="22" pageNumber="23">
<bibRefCitation id="B46B4B17FF97FF8015CFFBC8FEB9B22A" author="Kearn, G. C." box="[199,355,1062,1088]" pageId="22" pageNumber="23" pagination="301 - 311" refId="ref35977" refString="Kearn, G. C. (1970) The production, transfer and assimilation of spermatophores by Entobdella soleae, a monogenean skin parasite of the common sole. Parasitology, 60, 301 - 311." type="journal article" year="1970">Kearn (1970)</bibRefCitation>
reported that 
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<emphasis id="E28EEAF4FF97FF801700FBC8FDA9B255" box="[520,627,1062,1087]" italics="true" pageId="22" pageNumber="23">E. soleae</emphasis>
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typically possesses 2 seminal receptacles (
<figureCitation id="48C12A63FF97FF801163FBC8FB0DB22A" box="[1131,1239,1062,1088]" captionStart="FIGURE 37" captionStartId="32.[151,255,1897,1921]" captionTargetBox="[226,1392,197,1863]" captionTargetId="figure-76@32.[180,1424,146,1896]" captionTargetPageId="32" captionText="FIGURE 37. Entobdella soleae. A. Whole parasite in ventral view showing haptor papillae. Distribution of papillae derived from an SEM photograph by Dr J.A. Rees (see Kearn, 1998, fig. 4.3). B. Accessory sclerite enlarged. Abbreviation: cf, circular fibres associated with vaginal opening. Other abbreviations as in Figs. 26, 32. Scale bars: A, 1mm; B, 100µm." pageId="22" pageNumber="23">Fig. 37A</figureCitation>
), but individuals with 1, 3 and 4 receptacles occasionally occur. 
<bibRefCitation id="B46B4B17FF97FF8017D3FBA0FCA1B202" author="Kearn, G. C." box="[731,891,1102,1128]" pageId="22" pageNumber="23" pagination="301 - 311" refId="ref35977" refString="Kearn, G. C. (1970) The production, transfer and assimilation of spermatophores by Entobdella soleae, a monogenean skin parasite of the common sole. Parasitology, 60, 301 - 311." type="journal article" year="1970">Kearn (1970)</bibRefCitation>
found no evidence to indicate a relationship between receptacle number and age of parasites.
</paragraph>
<paragraph id="D04536E6FF97FF8015CFFB70FC55B0AA" blockId="22.[151,1436,1062,2008]" pageId="22" pageNumber="23">
The vaginal opening, diameter of 6 – 7, is located on the ventral surface (
<figureCitation id="48C12A63FF97FF801143FB70FB7CB2D2" box="[1099,1190,1182,1208]" captionStart="FIGURES 27 – 31" captionStartId="24.[151,269,1644,1668]" captionTargetBox="[264,1323,211,1585]" captionTargetId="figure-160@24.[264,1323,194,1585]" captionTargetPageId="24" captionText="FIGURES 27 – 31. Entobdella soleae. Light microscope photographs of toluidine-blue-stained serial resin sections through the vaginal region of an adult specimen. 27. Extended section showing the ventrally situated vaginal opening (arrow) in relation to the germarium (g). 28. Section through the narrow, distal entrance tube (et) of the vagina. 29. Section through the narrow, distal entrance tube (et) and 1 of the coils of the expanded proximal region (pr) of the vagina. 30. Section through the junction between the narrow entrance tube (et) and the expanded first coil of the proximal region (pr) of the vagina. 31. Extended section showing spacious coils of the proximal region (pr) of the vagina in relation to the vitelline reservoir (vr) and the germarium (g). Heavily stained vertical lines at right angles to the dorsal surface are folds in the resin section. Scale bars: 50µm." pageId="22" pageNumber="23">Fig. 27</figureCitation>
). The opening gives access to a relatively straight, narrow, distal tube (entrance tube), with a diameter between 3 and 8 (
<figureCitation id="48C12A63FF97FF80102EFB28FF62B362" captionStart="FIGURES 27 – 31" captionStartId="24.[151,269,1644,1668]" captionTargetBox="[264,1323,211,1585]" captionTargetId="figure-160@24.[264,1323,194,1585]" captionTargetPageId="24" captionText="FIGURES 27 – 31. Entobdella soleae. Light microscope photographs of toluidine-blue-stained serial resin sections through the vaginal region of an adult specimen. 27. Extended section showing the ventrally situated vaginal opening (arrow) in relation to the germarium (g). 28. Section through the narrow, distal entrance tube (et) of the vagina. 29. Section through the narrow, distal entrance tube (et) and 1 of the coils of the expanded proximal region (pr) of the vagina. 30. Section through the junction between the narrow entrance tube (et) and the expanded first coil of the proximal region (pr) of the vagina. 31. Extended section showing spacious coils of the proximal region (pr) of the vagina in relation to the vitelline reservoir (vr) and the germarium (g). Heavily stained vertical lines at right angles to the dorsal surface are folds in the resin section. Scale bars: 50µm." pageId="22" pageNumber="23">Figs. 28 – 30</figureCitation>
). In some sections through the distal end of the tube there is a distinct wall but this stained only lightly with toluidine blue (
<figureCitation id="48C12A63FF97FF801444FAF8FE7FB35A" box="[332,421,1302,1328]" captionStart="FIGURES 27 – 31" captionStartId="24.[151,269,1644,1668]" captionTargetBox="[264,1323,211,1585]" captionTargetId="figure-160@24.[264,1323,194,1585]" captionTargetPageId="24" captionText="FIGURES 27 – 31. Entobdella soleae. Light microscope photographs of toluidine-blue-stained serial resin sections through the vaginal region of an adult specimen. 27. Extended section showing the ventrally situated vaginal opening (arrow) in relation to the germarium (g). 28. Section through the narrow, distal entrance tube (et) of the vagina. 29. Section through the narrow, distal entrance tube (et) and 1 of the coils of the expanded proximal region (pr) of the vagina. 30. Section through the junction between the narrow entrance tube (et) and the expanded first coil of the proximal region (pr) of the vagina. 31. Extended section showing spacious coils of the proximal region (pr) of the vagina in relation to the vitelline reservoir (vr) and the germarium (g). Heavily stained vertical lines at right angles to the dorsal surface are folds in the resin section. Scale bars: 50µm." pageId="22" pageNumber="23">Fig. 28</figureCitation>
). There is no conspicuous heavily stained winding region similar to that between the entrance tube and the main vagina in “ 
<taxonomicName id="17FA4D65FF97FF801757FAD0FCDDB33D" authorityName="&quot; (Muller, 1776) Blainville" authorityYear="1818" baseAuthorityName="Muller" baseAuthorityYear="1776" box="[607,775,1342,1367]" class="Monogenea" family="Capsalidae" genus="Entobdella" kingdom="Animalia" order="Capsalidea" pageId="22" pageNumber="23" phylum="Platyhelminthes" rank="species" species="hippoglossi">
<emphasis id="E28EEAF4FF97FF801757FAD0FCDDB33D" box="[607,775,1342,1367]" italics="true" pageId="22" pageNumber="23">E. hippoglossi</emphasis>
</taxonomicName>
<typeStatus id="0F418844FF97FF801607FAD0FC98B332" box="[783,834,1342,1368]" pageId="22" pageNumber="23">type</typeStatus>
1” and “ 
<taxonomicName id="17FA4D65FF97FF8016A4FAD0FB8FB33D" authorityName="&quot; (Muller, 1776) Blainville" authorityYear="1818" baseAuthorityName="Muller" baseAuthorityYear="1776" box="[940,1109,1342,1367]" class="Monogenea" family="Capsalidae" genus="Entobdella" kingdom="Animalia" order="Capsalidea" pageId="22" pageNumber="23" phylum="Platyhelminthes" rank="species" species="hippoglossi">
<emphasis id="E28EEAF4FF97FF8016A4FAD0FB8FB33D" box="[940,1109,1342,1367]" italics="true" pageId="22" pageNumber="23">E. hippoglossi</emphasis>
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” from 
<taxonomicName id="17FA4D65FF97FF8011A3FAD0FA9EB33D" authorityName="Schmidt" authorityYear="1904" box="[1195,1348,1342,1367]" class="Actinopterygii" family="Pleuronectidae" genus="Hippoglossus" kingdom="Animalia" order="Pleuronectiformes" pageId="22" pageNumber="23" phylum="Chordata" rank="species" species="stenolepis">
<emphasis id="E28EEAF4FF97FF8011A3FAD0FA9EB33D" box="[1195,1348,1342,1367]" italics="true" pageId="22" pageNumber="23">H. stenolepis</emphasis>
</taxonomicName>
and no atrium close to the surface as in “ 
<taxonomicName id="17FA4D65FF97FF801714FA88FD1EB315" authorityName="&quot; (Muller, 1776) Blainville" authorityYear="1818" baseAuthorityName="Muller" baseAuthorityYear="1776" box="[540,708,1382,1407]" class="Monogenea" family="Capsalidae" genus="Entobdella" kingdom="Animalia" order="Capsalidea" pageId="22" pageNumber="23" phylum="Platyhelminthes" rank="species" species="hippoglossi">
<emphasis id="E28EEAF4FF97FF801714FA88FD1EB315" box="[540,708,1382,1407]" italics="true" pageId="22" pageNumber="23">E. hippoglossi</emphasis>
</taxonomicName>
<typeStatus id="0F418844FF97FF8017C3FA88FD27B3EA" box="[715,765,1382,1408]" pageId="22" pageNumber="23">type</typeStatus>
2”. Proximally, the distal tube of 
<taxonomicName id="17FA4D65FF97FF801180FA88FB29B315" baseAuthorityName="Van Beneden &amp; Hesse" baseAuthorityYear="1863" box="[1160,1267,1382,1407]" class="Monogenea" family="Capsalidae" genus="Entobdella" kingdom="Animalia" order="Capsalidea" pageId="22" pageNumber="23" phylum="Platyhelminthes" rank="species" species="soleae">
<emphasis id="E28EEAF4FF97FF801180FA88FB29B315" box="[1160,1267,1382,1407]" italics="true" pageId="22" pageNumber="23">E. soleae</emphasis>
</taxonomicName>
joins abruptly to the much more spacious convoluted proximal region of the vagina (
<figureCitation id="48C12A63FF97FF8016D3FA60FBEEB3C2" box="[987,1076,1422,1448]" captionStart="FIGURES 27 – 31" captionStartId="24.[151,269,1644,1668]" captionTargetBox="[264,1323,211,1585]" captionTargetId="figure-160@24.[264,1323,194,1585]" captionTargetPageId="24" captionText="FIGURES 27 – 31. Entobdella soleae. Light microscope photographs of toluidine-blue-stained serial resin sections through the vaginal region of an adult specimen. 27. Extended section showing the ventrally situated vaginal opening (arrow) in relation to the germarium (g). 28. Section through the narrow, distal entrance tube (et) of the vagina. 29. Section through the narrow, distal entrance tube (et) and 1 of the coils of the expanded proximal region (pr) of the vagina. 30. Section through the junction between the narrow entrance tube (et) and the expanded first coil of the proximal region (pr) of the vagina. 31. Extended section showing spacious coils of the proximal region (pr) of the vagina in relation to the vitelline reservoir (vr) and the germarium (g). Heavily stained vertical lines at right angles to the dorsal surface are folds in the resin section. Scale bars: 50µm." pageId="22" pageNumber="23">Fig. 30</figureCitation>
). Here the tube is narrow (~3) and has a thin wall with a stronger affinity for toluidine blue. The proximal coils of the vagina of 
<taxonomicName id="17FA4D65FF97FF801027FA58FA46B3A5" baseAuthorityName="Van Beneden &amp; Hesse" baseAuthorityYear="1863" box="[1327,1436,1462,1487]" class="Monogenea" family="Capsalidae" genus="Entobdella" kingdom="Animalia" order="Capsalidea" pageId="22" pageNumber="23" phylum="Platyhelminthes" rank="species" species="soleae">
<emphasis id="E28EEAF4FF97FF801027FA58FA46B3A5" box="[1327,1436,1462,1487]" italics="true" pageId="22" pageNumber="23">E. soleae</emphasis>
</taxonomicName>
contain stored sperm and are more spacious than those of “ 
<taxonomicName id="17FA4D65FF97FF801642FA30FC28B39D" authorityName="&quot; (Muller, 1776) Blainville" authorityYear="1818" baseAuthorityName="Muller" baseAuthorityYear="1776" box="[842,1010,1502,1527]" class="Monogenea" family="Capsalidae" genus="Entobdella" kingdom="Animalia" order="Capsalidea" pageId="22" pageNumber="23" phylum="Platyhelminthes" rank="species" species="hippoglossi">
<emphasis id="E28EEAF4FF97FF801642FA30FC28B39D" box="[842,1010,1502,1527]" italics="true" pageId="22" pageNumber="23">E. hippoglossi</emphasis>
</taxonomicName>
<typeStatus id="0F418844FF97FF8016F1FA30FBEDB392" box="[1017,1079,1502,1528]" pageId="22" pageNumber="23">types</typeStatus>
1 and 2” and “ 
<taxonomicName id="17FA4D65FF97FF8011EFFA30FA54B39D" authorityName="&quot; (Muller, 1776) Blainville" authorityYear="1818" baseAuthorityName="Muller" baseAuthorityYear="1776" box="[1255,1422,1502,1527]" class="Monogenea" family="Capsalidae" genus="Entobdella" kingdom="Animalia" order="Capsalidea" pageId="22" pageNumber="23" phylum="Platyhelminthes" rank="species" species="hippoglossi">
<emphasis id="E28EEAF4FF97FF8011EFFA30FA54B39D" box="[1255,1422,1502,1527]" italics="true" pageId="22" pageNumber="23">E. hippoglossi</emphasis>
</taxonomicName>
” from 
<taxonomicName id="17FA4D65FF97FF8015D3F9E8FEA2B075" authorityName="Schmidt" authorityYear="1904" box="[219,376,1542,1567]" class="Actinopterygii" family="Pleuronectidae" genus="Hippoglossus" kingdom="Animalia" order="Pleuronectiformes" pageId="22" pageNumber="23" phylum="Chordata" rank="species" species="stenolepis">
<emphasis id="E28EEAF4FF97FF8015D3F9E8FEA2B075" box="[219,376,1542,1567]" italics="true" pageId="22" pageNumber="23">H. stenolepis</emphasis>
</taxonomicName>
, being arranged in a compact entangled knot (
<figureCitation id="48C12A63FF97FF8016BDF9E8FB91B04A" box="[949,1099,1542,1568]" captionStart="FIGURES 27 – 31" captionStartId="24.[151,269,1644,1668]" captionTargetBox="[264,1323,211,1585]" captionTargetId="figure-160@24.[264,1323,194,1585]" captionTargetPageId="24" captionText="FIGURES 27 – 31. Entobdella soleae. Light microscope photographs of toluidine-blue-stained serial resin sections through the vaginal region of an adult specimen. 27. Extended section showing the ventrally situated vaginal opening (arrow) in relation to the germarium (g). 28. Section through the narrow, distal entrance tube (et) of the vagina. 29. Section through the narrow, distal entrance tube (et) and 1 of the coils of the expanded proximal region (pr) of the vagina. 30. Section through the junction between the narrow entrance tube (et) and the expanded first coil of the proximal region (pr) of the vagina. 31. Extended section showing spacious coils of the proximal region (pr) of the vagina in relation to the vitelline reservoir (vr) and the germarium (g). Heavily stained vertical lines at right angles to the dorsal surface are folds in the resin section. Scale bars: 50µm." pageId="22" pageNumber="23">Figs. 30, 31</figureCitation>
). The proximal coils of the vagina lie close to the wall of the vitelline reservoir (
<bibRefCitation id="B46B4B17FF97FF801609F9C0FC4AB022" author="Kearn, G. C." box="[769,912,1582,1608]" pageId="22" pageNumber="23" pagination="301 - 311" refId="ref35977" refString="Kearn, G. C. (1970) The production, transfer and assimilation of spermatophores by Entobdella soleae, a monogenean skin parasite of the common sole. Parasitology, 60, 301 - 311." type="journal article" year="1970">Kearn 1970</bibRefCitation>
), but the junction with this reservoir was not seen in resin sections. In a live adult specimen with anterior hamulus length of about 570 (total body length about 5500), the distal narrow tube is about 95 long, while the distance measured in a straight line between the proximal end of this tube and the vitelline reservoir is about 265.
</paragraph>
<paragraph id="D04536E6FF97FF8015CFF920FD9FB1B2" blockId="22.[151,1436,1062,2008]" pageId="22" pageNumber="23">
Each anterior adhesive pad of 
<taxonomicName id="17FA4D65FF97FF80173EF920FD7EB08D" baseAuthorityName="Van Beneden &amp; Hesse" baseAuthorityYear="1863" box="[566,676,1742,1767]" class="Monogenea" family="Capsalidae" genus="Entobdella" kingdom="Animalia" order="Capsalidea" pageId="22" pageNumber="23" phylum="Platyhelminthes" rank="species" species="soleae">
<emphasis id="E28EEAF4FF97FF80173EF920FD7EB08D" box="[566,676,1742,1767]" italics="true" pageId="22" pageNumber="23">E. soleae</emphasis>
</taxonomicName>
is subdivided into 3 areas (
<bibRefCitation id="B46B4B17FF97FF8016FAF920FA81B082" author="Kearn, G. C. &amp; Evans-Gowing, R." box="[1010,1371,1742,1768]" pageId="22" pageNumber="23" pagination="1583 - 1593" refId="ref36360" refString="Kearn, G. C. &amp; Evans-Gowing, R. (1998) Attachment and detachment of the anterior adhesive pads of the monogenean (platyhelminth) parasite Entobdella soleae from the skin of the common sole (Solea solea). International Journal for Parasitology, 28, 1583 - 1593." type="journal article" year="1998">Kearn &amp; Evans-Gowing 1998</bibRefCitation>
) (see also 
<figureCitation id="48C12A63FF97FF8015C7F918FEE4B17A" box="[207,318,1782,1808]" captionStart="FIGURE 37" captionStartId="32.[151,255,1897,1921]" captionTargetBox="[226,1392,197,1863]" captionTargetId="figure-76@32.[180,1424,146,1896]" captionTargetPageId="32" captionText="FIGURE 37. Entobdella soleae. A. Whole parasite in ventral view showing haptor papillae. Distribution of papillae derived from an SEM photograph by Dr J.A. Rees (see Kearn, 1998, fig. 4.3). B. Accessory sclerite enlarged. Abbreviation: cf, circular fibres associated with vaginal opening. Other abbreviations as in Figs. 26, 32. Scale bars: A, 1mm; B, 100µm." pageId="22" pageNumber="23">Fig. 37A</figureCitation>
). The accessory sclerites are strongly curved (hooked) (
<bibRefCitation id="B46B4B17FF97FF8016D1F918FBB0B17A" author="Kearn, G. C." box="[985,1130,1782,1808]" pageId="22" pageNumber="23" pagination="327 - 335" refId="ref35883" 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>
; 
<figureCitation id="48C12A63FF97FF801170F918FB00B17A" box="[1144,1242,1782,1808]" captionStart="FIGURE 15" captionStartId="15.[151,255,1634,1658]" captionTargetBox="[215,1390,834,1594]" captionTargetId="figure-365@15.[184,1411,794,1641]" captionTargetPageId="15" captionText="FIGURE 15. Entobdella spp. Accessory sclerites of adults. A. “E. hippoglossi type 1” from H. hippoglossus. B. “E. hippoglossi type 2” from H. hippoglossus. C. “Entobdella hippoglossi” from H. stenolepis. D. E. steingroeveri. E. E. squamula from Paralichthys californicus. F, G. accessory sclerites from 2 individuals of different sizes (ages) of a proposed new species from Hippoglossina macrops (F, Entobdella sp. 1 from Dr. M.T. González; G, from LUCE; see Table 2). H. E. brattstroemi from P. adspersus (Entobdella sp. 2 from Dr M.T. González; Table 2). J. E. soleae from Solea solea. K. E. rosaceus from Zalembius rosaceus. All sclerites drawn to the same scale with the distal region uppermost. Scale bars: 100µm." pageId="22" pageNumber="23">Fig. 15J</figureCitation>
) and the tendon associated with each terminates on the anterior (proximal) end of the anterior hamulus (
<figureCitation id="48C12A63FF97FF8011A9F8F0FACAB152" box="[1185,1296,1822,1848]" captionStart="FIGURE 37" captionStartId="32.[151,255,1897,1921]" captionTargetBox="[226,1392,197,1863]" captionTargetId="figure-76@32.[180,1424,146,1896]" captionTargetPageId="32" captionText="FIGURE 37. Entobdella soleae. A. Whole parasite in ventral view showing haptor papillae. Distribution of papillae derived from an SEM photograph by Dr J.A. Rees (see Kearn, 1998, fig. 4.3). B. Accessory sclerite enlarged. Abbreviation: cf, circular fibres associated with vaginal opening. Other abbreviations as in Figs. 26, 32. Scale bars: A, 1mm; B, 100µm." pageId="22" pageNumber="23">Fig. 37A</figureCitation>
). The entire haptor ventral surface is covered with papillae and papillae lateral to the median sclerites are arranged in radiating rows (see Kearn 1998, fig. 4.3 and 
<figureCitation id="48C12A63FF97FF801778F880FD07B1E2" box="[624,733,1902,1928]" captionStart="FIGURE 37" captionStartId="32.[151,255,1897,1921]" captionTargetBox="[226,1392,197,1863]" captionTargetId="figure-76@32.[180,1424,146,1896]" captionTargetPageId="32" captionText="FIGURE 37. Entobdella soleae. A. Whole parasite in ventral view showing haptor papillae. Distribution of papillae derived from an SEM photograph by Dr J.A. Rees (see Kearn, 1998, fig. 4.3). B. Accessory sclerite enlarged. Abbreviation: cf, circular fibres associated with vaginal opening. Other abbreviations as in Figs. 26, 32. Scale bars: A, 1mm; B, 100µm." pageId="22" pageNumber="23">Fig. 37A</figureCitation>
). 
<bibRefCitation id="B46B4B17FF97FF8017F8F880FC57B1E2" author="Lyons, K. M." box="[752,909,1902,1928]" pageId="22" pageNumber="23" pagination="471 - 480" refId="ref36639" refString="Lyons, K. M. (1973) Scanning and transmission electron microscope studies on the sensory sucker papillae of the fish parasite Entobdella soleae (Monogenea). Zeitschrift fur Zellforschung, 137, 471 - 480." type="journal article" year="1973">Lyons (1973)</bibRefCitation>
estimated that there are more than 800 papillae on the haptor ventral surface. The eyes are conspicuous in the adult and glands of Goto are present (
<bibRefCitation id="B46B4B17FF97FF80105AF878FF3BB1B2" author="Kearn, G. C." pageId="22" pageNumber="23" pagination="161 - 187" refId="ref36015" refString="Kearn, G. C. (1971 a) The physiology and behaviour of the monogenean skin parasite Entobdella soleae in relation to its host (Solea solea). In: Fallis, A. M. (Ed.). Ecology and Physiology of Parasites, a Symposium. University of Toronto Press, Toronto, pp. 161 - 187." type="book chapter" year="1971">Kearn 1971a</bibRefCitation>
, fig. 2; see also our 
<figureCitation id="48C12A63FF97FF8014C4F850FDE3B1B2" box="[460,569,1982,2008]" captionStart="FIGURE 37" captionStartId="32.[151,255,1897,1921]" captionTargetBox="[226,1392,197,1863]" captionTargetId="figure-76@32.[180,1424,146,1896]" captionTargetPageId="32" captionText="FIGURE 37. Entobdella soleae. A. Whole parasite in ventral view showing haptor papillae. Distribution of papillae derived from an SEM photograph by Dr J.A. Rees (see Kearn, 1998, fig. 4.3). B. Accessory sclerite enlarged. Abbreviation: cf, circular fibres associated with vaginal opening. Other abbreviations as in Figs. 26, 32. Scale bars: A, 1mm; B, 100µm." pageId="22" pageNumber="23">Fig. 37A</figureCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>