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<document id="90DEE4FF8604A2B80D5A33FE106336F8" ID-DOI="10.1046/j.1096-3642.2003.00058.x" ID-ISSN="0024-4082" ID-Zenodo-Dep="5437607" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.taxonomicNames_approvedBy="felipe" checkinTime="1630437499139" checkinUser="carolina" docAuthor="Strong, Ellen E." docDate="2003" docId="03B6B923EE15FFC78E4C675239A06ADE" docLanguage="en" docName="j.1096-3642.2003.00058.x.pdf" docOrigin="Zoological Journal of the Linnean Society 137 (4)" docSource="https://academic.oup.com/zoolinnean/article-lookup/doi/10.1046/j.1096-3642.2003.00058.x" docStyle="DocumentStyle:0DD8C314D74634CE09062A86991413F8.2:ZoolJLinnSoc.2002-2009.journal_article" docStyleId="0DD8C314D74634CE09062A86991413F8" docStyleName="ZoolJLinnSoc.2002-2009.journal_article" docStyleVersion="2" docTitle="Ilyanassa obsoletus" docType="treatment" docVersion="3" lastPageNumber="484" masterDocId="FF8FC15BEE36FFE28F4B66213B1A6964" masterDocTitle="Refining molluscan characters: morphology, character coding and a phylogeny of the Caenogastropoda" masterLastPageNumber="554" masterPageNumber="447" pageNumber="482" updateTime="1699271261741" updateUser="plazi" zenodo-license-document="CC-BY-4.0" zenodo-license-figures="CC-BY-4.0">
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<mods:title id="BF9612B7650002AC2E63473E2885977D">Refining molluscan characters: morphology, character coding and a phylogeny of the Caenogastropoda</mods:title>
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<mods:namePart id="81BB0413486A17B41EB7409DE363C23D">Strong, Ellen E.</mods:namePart>
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<mods:title id="A9BF4EFE6E8BFEAD833867CB67135E0D">Zoological Journal of the Linnean Society</mods:title>
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<mods:date id="BB47004A8B95126542C356EE25C5D0D7">2003</mods:date>
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<mods:number id="EC0F57160E6B220DE0AB2C962068664A">2003-04-30</mods:number>
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<treatment id="03B6B923EE15FFC78E4C675239A06ADE" LSID="urn:lsid:plazi:treatment:03B6B923EE15FFC78E4C675239A06ADE" httpUri="http://treatment.plazi.org/id/03B6B923EE15FFC78E4C675239A06ADE" lastPageId="37" lastPageNumber="484" pageId="35" pageNumber="482">
<subSubSection id="C3055BBEEE15FFC18E4C6752398C68EE" box="[263,662,370,394]" pageId="35" pageNumber="482" type="nomenclature">
<paragraph id="8BA00835EE15FFC18E4C6752398C68EE" blockId="35.[263,662,338,394]" box="[263,662,370,394]" pageId="35" pageNumber="482">
<heading id="D0E8BF59EE15FFC18E4C6752398C68EE" box="[263,662,370,394]" centered="true" fontSize="9" level="2" pageId="35" pageNumber="482" reason="2">
<taxonomicName id="4C1F73B6EE15FFC18E4C6752398C68EE" ID-CoL="84XLS" authority="(SAY 1822)" baseAuthorityName="SAY" baseAuthorityYear="1822" box="[263,662,370,394]" class="Gastropoda" family="Nassariidae" genus="Ilyanassa" higherTaxonomySource="GBIF" kingdom="Animalia" order="Neogastropoda" pageId="35" pageNumber="482" phylum="Mollusca" rank="species" species="obsoletus">
<emphasis id="B96BD427EE15FFC18E4C6752390A68EC" box="[263,528,371,394]" italics="true" pageId="35" pageNumber="482">ILYANASSA OBSOLETUS</emphasis>
(SAY 1822)
</taxonomicName>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="C3055BBEEE15FFC78FD067BB39A06ADE" lastPageId="37" lastPageNumber="484" pageId="35" pageNumber="482" type="materials_examined">
<paragraph id="8BA00835EE15FFC18FD067BB3A7668CB" blockId="35.[155,656,410,463]" box="[155,364,410,431]" pageId="35" pageNumber="482">
<emphasis id="B96BD427EE15FFC18FD067BB3A7668CB" box="[155,364,410,431]" italics="true" pageId="35" pageNumber="482">Material examined</emphasis>
</paragraph>
<paragraph id="8BA00835EE15FFC18FD06798398A68AB" blockId="35.[155,656,410,463]" box="[155,656,441,463]" pageId="35" pageNumber="482">
Cape Henlopen,
<collectingRegion id="49DBC6D7EE15FFC18E1267983ADC68AB" box="[345,454,441,463]" country="United States of America" name="Delaware" pageId="35" pageNumber="482">Delaware</collectingRegion>
(
<collectionCode id="ED0E90F0EE15FFC18E9E6798393F68AA" box="[469,549,441,462]" country="USA" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34871" lsid="urn:lsid:biocol.org:col:34871" name="Smithsonian Institution, National Museum of Natural History" pageId="35" pageNumber="482" type="Museum">USNM</collectionCode>
890946).
</paragraph>
<paragraph id="8BA00835EE15FFC18FD0642039296B73" blockId="35.[155,770,513,719]" box="[155,563,513,535]" pageId="35" pageNumber="482">
<emphasis id="B96BD427EE15FFC18FD0642039296B73" box="[155,563,513,535]" italics="true" pageId="35" pageNumber="482">External anatomy and mantle cavity</emphasis>
</paragraph>
<paragraph id="8BA00835EE15FFC18FD064013AA26BAB" blockId="35.[155,770,513,719]" pageId="35" pageNumber="482">
Mantle edge smooth. Siphon long with basal flaps (
<figureCitation id="132414B0EE15FFC18FE9641E3BEF6B30" box="[162,245,575,597]" captionStart="Figure 7" captionStartId="51.[155,233,1525,1544]" captionTargetBox="[373,1221,197,1490]" captionTargetId="figure-270@51.[363,1227,197,1493]" captionTargetPageId="51" captionText="Figure 7. Morphology of cephalic haemocoel among derived caenogastropods. (A) Panarona clavatula. Siphon obscured by left cephalic tentacle and is not visible in this view. Note large, modified left cephalic tentacle and large proboscis retractor connecting to columellar muscle. (B) Ilyanassa obsoletus. Note second visceral ganglion lying below pericardium and large radular retractors connecting to base of proboscis sheath; ampulla of gland of Leiblein visible within afferent renal vessel. Duct of gland of Leiblein stippled. (C) Nitidiscala tinctum. Note compartmentalized tissue supporting jaw and buccal ganglia lying between salivary glands where the latter enter buccal cavity. (D) Prunum apicinum. Note duct of poison gland opening mid-ventrally to base of buccal cavity, just behind opening of accessory salivary gland. (E) Conus jaspideus. Siphon not visible in this view. Note long coiled duct of poison gland; buccal ganglion visible where short arm of radular sac enters buccal cavity. Base of proboscis sheath is shown, but rhynchocoel has not been opened. Scale bar = 1 mm. Abbreviations: asg, accessory salivary gland; dasg, duct of accessory salivary gland; e, oesophagus; ec, oesophageal caecum; gl, gland of Leiblein; j, jaw; kd, kidney; nr, circum-oesophageal nerve ring; os, osphradium; per, pericardium; ps, proboscis sheath; rs, long arm of radular sac; sb, sub-oesophageal ganglion; sgd, salivary gland duct; sgl, salivary glands; si, siphon; sig, siphonal ganglion; sp, supra-oesophageal ganglion; tn, tentacular nerve; vg, visceral ganglion; vl, valve of Leiblein; z, zygoneury." figureDoi="http://doi.org/10.5281/zenodo.5437621" httpUri="https://zenodo.org/record/5437621/files/figure.png" pageId="35" pageNumber="482">Fig. 7B</figureCitation>
, si), opening to bipectinate osphradium (os). Anterior pedal gland opening under thick propodial flap. Ventral pedal gland present in females, opening mid-ventrally via small slit, just behind front edge of foot. Operculum present.
</paragraph>
<paragraph id="8BA00835EE15FFC18FD065203A676A73" blockId="35.[155,771,769,1773]" box="[155,381,769,791]" pageId="35" pageNumber="482">
<emphasis id="B96BD427EE15FFC18FD065203A676A73" box="[155,381,769,791]" italics="true" pageId="35" pageNumber="482">Reproductive system</emphasis>
</paragraph>
<paragraph id="8BA00835EE15FFC18FD0650139A26F89" blockId="35.[155,771,769,1773]" pageId="35" pageNumber="482">
Gonad dorsally overlying digestive gland on right, to base of gastric chamber. Gonopericardial canal present behind base of mantle cavity (
<figureCitation id="132414B0EE15FFC18D3B657C39A16A16" box="[624,699,861,883]" captionStart="Figure 2" captionStartId="42.[142,220,1619,1638]" captionTargetBox="[319,1244,198,1585]" captionTargetId="figure-210@42.[319,1245,198,1587]" captionTargetPageId="42" captionText="Figure 2. Female reproductive morphology. (A) Lampanella minima. Note connective tissue between renal oviduct and pericardium. (B) Crepidula plana. (C) Petaloconchus varians. (D) Strombus mutabilis. Note egg groove from capsule gland down side of foot. Dotted line indicates double-groove portion of albumen gland. (E) Cypraea robertsi. Note duct of receptacle at posterior end of albumen gland. (F) Panarona clavatula. (G) Prunum apicinum. Note that base of mantle cavity lies between the capsule and albumen glands. (H) Urosalpinx cinerea. Ventro-lateral lobe of capsule gland is shown by transparency. Note blind diverticulum off renal oviduct and small aperture connecting vestibule and bursa copulatrix. Base of mantle cavity indicated by arrow. (I) Ilyanassa obsoletus. Note shallow pocket in distal end of capsule gland, perhaps functioning as bursa. Scale bar = 1 mm. Abbreviations: ag, albumen gland; bc, bursa copulatrix; cg, capsule gland; fo, female opening; gpc, gonopericardial canal; igl, ingesting gland; mc, base of mantle cavity; per, pericardium; rcs, seminal receptacle; spb, spermatophore bursa; u, uterine ball; v, vagina." figureDoi="http://doi.org/10.5281/zenodo.5437611" httpUri="https://zenodo.org/record/5437611/files/figure.png" pageId="35" pageNumber="482">Fig. 2I</figureCitation>
, gpc); small flap marking entrance of canal to pericardium. Oviduct opening to small albumen gland with Ushaped lumen (ag). Three to four small pouches along crest of albumen gland possibly functioning in storage of sperm; no sperm present. Albumen gland narrowing and opening to ventral channel of capsule gland (cg), dorsally giving off narrow duct leading to lobate ingesting gland (igl). No sperm present in duct of ingesting gland. Capsule gland comprising paired, glandular laminae bearing regionated epithelium, enclosing tall, narrow lumen. Proximal capsule gland bordering albumen gland turning briefly to left, then turning anteriorly and continuing towards female opening. Ventral channel opening at anterior end of capsule gland, to short, narrow vagina, terminating in simple female opening (fo). Muscular bursa and vestibule absent; glandular tissue of distal capsule gland forming shallow pocket, possibly functioning as bursa. Vas deferens emerging from testis and expanding into large, convoluted seminal vesicle. Vas deferens narrowing near back wall of kidney. Several strands of connective tissue connecting vas deferens and pericardial wall; gonopericardial canal absent. Vas deferens opening to narrow, convoluted prostate at base of mantle cavity (
<figureCitation id="132414B0EE15FFC18E58607D3A726F16" box="[275,360,1628,1650]" captionStart="Figure 3" captionStartId="44.[142,220,1712,1731]" captionTargetBox="[277,1288,197,1677]" captionTargetId="figure-151@44.[277,1288,166,1683]" captionTargetPageId="44" captionText="Figure 3. Male reproductive morphology. (A) Lampanella minima. Note connective tissue between renal vas deferens and pericardium and accessory lobe of prostatic tissue on mantle cavity floor alongside proximal prostate. (B) Petaloconchus varians. (C) Crepidula plana. Note terminal papilla. (D) Strombus mutabilis. Note seminal groove continuous along ventral aspect of penis. (E) Cypraea robertsi. Note seminal groove continuous along ventral aspect of penis. (F) Prunum apicinum. Penial duct not shown. Arrow indicates slit-like communication between proximal prostate and mantle cavity. (G) Ilyanassa obsoletus. Arrow indicates fused, tube-like communication between proximal prostate and mantle cavity. Scale bar = 1 mm. Abbreviations: ped, penial duct; pe, penis; per, pericardium; pr, prostate; pvd, pallial vas deferens; sg, seminal groove." figureDoi="http://doi.org/10.5281/zenodo.5437613" httpUri="https://zenodo.org/record/5437613/files/figure.png" pageId="35" pageNumber="482">Fig. 3G</figureCitation>
). Small diverticulum from proximal prostate ending blindly. Pallial vas deferens crossing neck and entering base of penis (pe). Penis long and dorso-ventrally flattened with spatulate tip. Penial duct (ped) running subcentrally to tip of penis.
</paragraph>
<paragraph id="8BA00835EE15FFC18FD0613C3A746E56" blockId="35.[155,770,1821,1905]" box="[155,366,1821,1842]" pageId="35" pageNumber="482">
<emphasis id="B96BD427EE15FFC18FD0613C3A746E56" box="[155,366,1821,1842]" italics="true" pageId="35" pageNumber="482">Alimentary System</emphasis>
</paragraph>
<paragraph id="8BA00835EE15FFC18FD0611C3FB268F5" blockId="35.[155,770,1821,1905]" lastBlockId="35.[819,1435,196,1291]" pageId="35" pageNumber="482">
<emphasis id="B96BD427EE15FFC18FD0611C3BE36E36" box="[155,249,1853,1874]" italics="true" pageId="35" pageNumber="482">Foregut.</emphasis>
Salivary gland ducts (
<figureCitation id="132414B0EE15FFC18D67611D39656E35" box="[556,639,1852,1874]" captionStart="Figure 7" captionStartId="51.[155,233,1525,1544]" captionTargetBox="[373,1221,197,1490]" captionTargetId="figure-270@51.[363,1227,197,1493]" captionTargetPageId="51" captionText="Figure 7. Morphology of cephalic haemocoel among derived caenogastropods. (A) Panarona clavatula. Siphon obscured by left cephalic tentacle and is not visible in this view. Note large, modified left cephalic tentacle and large proboscis retractor connecting to columellar muscle. (B) Ilyanassa obsoletus. Note second visceral ganglion lying below pericardium and large radular retractors connecting to base of proboscis sheath; ampulla of gland of Leiblein visible within afferent renal vessel. Duct of gland of Leiblein stippled. (C) Nitidiscala tinctum. Note compartmentalized tissue supporting jaw and buccal ganglia lying between salivary glands where the latter enter buccal cavity. (D) Prunum apicinum. Note duct of poison gland opening mid-ventrally to base of buccal cavity, just behind opening of accessory salivary gland. (E) Conus jaspideus. Siphon not visible in this view. Note long coiled duct of poison gland; buccal ganglion visible where short arm of radular sac enters buccal cavity. Base of proboscis sheath is shown, but rhynchocoel has not been opened. Scale bar = 1 mm. Abbreviations: asg, accessory salivary gland; dasg, duct of accessory salivary gland; e, oesophagus; ec, oesophageal caecum; gl, gland of Leiblein; j, jaw; kd, kidney; nr, circum-oesophageal nerve ring; os, osphradium; per, pericardium; ps, proboscis sheath; rs, long arm of radular sac; sb, sub-oesophageal ganglion; sgd, salivary gland duct; sgl, salivary glands; si, siphon; sig, siphonal ganglion; sp, supra-oesophageal ganglion; tn, tentacular nerve; vg, visceral ganglion; vl, valve of Leiblein; z, zygoneury." figureDoi="http://doi.org/10.5281/zenodo.5437621" httpUri="https://zenodo.org/record/5437621/files/figure.png" pageId="35" pageNumber="482">Fig. 7B</figureCitation>
, sgd) bypassing nerve ring, embedded anteriorly within dorsal folds, then curving around base of buccal cavity, opening ventrally under elongate odontophore. Accessory salivary glands and jaws absent. Three nerves from each buccal ganglion and each cerebral ganglion innervating large pleurembolic proboscis. Nerves and buccal artery entering proboscis ventrally, passing through proboscis basal septum.
</paragraph>
<paragraph id="8BA00835EE15FFC18C0067BA38B76D72" blockId="35.[819,1435,196,1291]" pageId="35" pageNumber="482">Paired cartilages fused anteriorly. Lateral edges of cartilages curving dorsally, forming gutter supporting radular ribbon. Deep sulcus dorsally enclosing rachiglossate radula. Odontophore long, narrow, and dorsoventrally flattened. Radula passing over bending plane of anterior tip, extending posteriorly into deep sublingual cavity. Subradular organ absent. Unfused posterior tips of cartilages, becoming enveloped in thick sheath of muscle fibres, flanking long, straight radular sac. Paired retractor muscles connecting each bolster tip and radular sac to ventral proboscis wall (ps). Thick muscular sheet dorsally connecting bolsters. Two large sets of retractors, connecting inner sides of bolsters to dorso-lateral proboscis walls. Single, unpaired set of retractors attaching ventrally to buccal mass and posterior proboscis sheath. Two large protractors attaching laterally to posterior ends of bolsters and anteriorly to sides of proboscis. Large buccal mass filling cephalic haemocoel when retracted, projecting well past posterior end of proboscis sheath.</paragraph>
<paragraph id="8BA00835EE15FFC18C00623E3E656C6F" blockId="35.[819,1435,196,1291]" pageId="35" pageNumber="482">
Dorsal folds beginning just behind mouth, continuing into anterior oesophagus, gradually migrating ventrally. Ventral folds absent within anterior most oesophagus (
<figureCitation id="132414B0EE15FFC18C8E625A3F3F6DF5" box="[965,1061,1147,1169]" captionStart="Figure 10" captionStartId="56.[142,220,1725,1744]" captionTargetBox="[283,1280,197,1693]" captionTargetId="figure-143@56.[283,1281,197,1693]" captionTargetPageId="56" captionText="Figure 10. Histology of anterior oesophagus; ventral folding absent. (A) Neverita duplicata. Scale bar = 0.5 mm. (B) Nitidiscala tinctum. Scale bar = 0.5 mm. (C) Panarona clavatula. Scale bar = 0.5 mm. (D) Prunum apicinum. Scale bar = 0.5 mm. (E) Conus jaspideus. Scale bar = 0.5 mm. (F) Ilyanassa obsoletus. Scale bar = 0.5 mm. (G) Urosalpinx cinerea. Scale bar = 1 mm. Abbreviations: asg, accessory salivary gland; ca, radular cartilage; ceg, cerebral ganglion; df, dorsal fold; dl, duct of gland of Leiblein; e, oesophagus; int, intestine; j, jaw; ps, proboscis sheath; r, radula; sgd, salivary gland duct; sgl, salivary gland; vc, ventral channel." figureDoi="http://doi.org/10.5281/zenodo.5437627" httpUri="https://zenodo.org/record/5437627/files/figure.png" pageId="35" pageNumber="482">Fig. 10F</figureCitation>
). Walls of distal anterior oesophagus becoming uniformly highly folded. Valve of Leiblein (
<figureCitation id="132414B0EE15FFC18CE9629938EF6DA9" box="[930,1013,1208,1230]" captionStart="Figure 7" captionStartId="51.[155,233,1525,1544]" captionTargetBox="[373,1221,197,1490]" captionTargetId="figure-270@51.[363,1227,197,1493]" captionTargetPageId="51" captionText="Figure 7. Morphology of cephalic haemocoel among derived caenogastropods. (A) Panarona clavatula. Siphon obscured by left cephalic tentacle and is not visible in this view. Note large, modified left cephalic tentacle and large proboscis retractor connecting to columellar muscle. (B) Ilyanassa obsoletus. Note second visceral ganglion lying below pericardium and large radular retractors connecting to base of proboscis sheath; ampulla of gland of Leiblein visible within afferent renal vessel. Duct of gland of Leiblein stippled. (C) Nitidiscala tinctum. Note compartmentalized tissue supporting jaw and buccal ganglia lying between salivary glands where the latter enter buccal cavity. (D) Prunum apicinum. Note duct of poison gland opening mid-ventrally to base of buccal cavity, just behind opening of accessory salivary gland. (E) Conus jaspideus. Siphon not visible in this view. Note long coiled duct of poison gland; buccal ganglion visible where short arm of radular sac enters buccal cavity. Base of proboscis sheath is shown, but rhynchocoel has not been opened. Scale bar = 1 mm. Abbreviations: asg, accessory salivary gland; dasg, duct of accessory salivary gland; e, oesophagus; ec, oesophageal caecum; gl, gland of Leiblein; j, jaw; kd, kidney; nr, circum-oesophageal nerve ring; os, osphradium; per, pericardium; ps, proboscis sheath; rs, long arm of radular sac; sb, sub-oesophageal ganglion; sgd, salivary gland duct; sgl, salivary glands; si, siphon; sig, siphonal ganglion; sp, supra-oesophageal ganglion; tn, tentacular nerve; vg, visceral ganglion; vl, valve of Leiblein; z, zygoneury." figureDoi="http://doi.org/10.5281/zenodo.5437621" httpUri="https://zenodo.org/record/5437621/files/figure.png" pageId="35" pageNumber="482">Fig. 7B</figureCitation>
, vl) present proximal to nerve ring; gland of Leiblein present with terminal ampulla. Dorsal folds weakly developed within mid-oesophagus.
</paragraph>
<paragraph id="8BA00835EE15FFC18C7863123E976E14" blockId="35.[819,1435,1331,1905]" pageId="35" pageNumber="482">
<emphasis id="B96BD427EE15FFC18C78631238946C2C" box="[819,910,1331,1352]" italics="true" pageId="35" pageNumber="482">Midgut.</emphasis>
Oesophagus opening ventrally on left (
<figureCitation id="132414B0EE15FFC18C71637038876C03" box="[826,925,1361,1383]" captionStart="Figure 19" captionStartId="69.[155,233,1787,1806]" captionTargetBox="[276,1305,197,1754]" captionTargetId="figure-91@69.[276,1315,197,1754]" captionTargetPageId="69" captionText="Figure 19. Midgut morphology. White arrows indicate direction of ciliary currents. (A) Ilyanassa obsoletus, dorsal view. (B) Urosalpinx cinerea, dorsal view. Scale bar = 1 mm. Abbreviations: ce, caecal extension; dgd, digestive gland duct; e, oesophagus; gp, glandular pad; gs, gastric shield; ig, intestinal groove; int, intestine; po, pouch; sa, sorting area; ss, style sac; t1, major typhlosole; t2, minor typhlosole." figureDoi="http://doi.org/10.5281/zenodo.5437647" httpUri="https://zenodo.org/record/5437647/files/figure.png" pageId="35" pageNumber="482">Fig. 19A</figureCitation>
). Well-developed sorting area present to left of oesophagus (sa). Paired digestive gland ducts (dgd) opening to right of oesophagus. Long, narrow glandular fold (gp), separating digestive gland ducts from small, ventro-lateral gastric shield (gs). Posterior gastric chamber extremely elongate (ce), lined with textured, knobby epithelium roughly arranged in longitudinal folds; irregular tufts of cilia crowning crests of folds. Ciliary currents within gastric chamber flowing posteriorly on left and anteriorly on right. Ciliary currents flowing clockwise in sorting area and adjacent to intestinal groove. Large, well-developed style sac region bearing two large typhlosoles, each possessing ciliated strip of raised cilia along inner (style sac) margins (
<figureCitation id="132414B0EE15FFC18B5E60DE3F646E70" box="[1045,1150,1791,1813]" captionStart="Figure 19" captionStartId="69.[155,233,1787,1806]" captionTargetBox="[276,1305,197,1754]" captionTargetId="figure-91@69.[276,1315,197,1754]" captionTargetPageId="69" captionText="Figure 19. Midgut morphology. White arrows indicate direction of ciliary currents. (A) Ilyanassa obsoletus, dorsal view. (B) Urosalpinx cinerea, dorsal view. Scale bar = 1 mm. Abbreviations: ce, caecal extension; dgd, digestive gland duct; e, oesophagus; gp, glandular pad; gs, gastric shield; ig, intestinal groove; int, intestine; po, pouch; sa, sorting area; ss, style sac; t1, major typhlosole; t2, minor typhlosole." figureDoi="http://doi.org/10.5281/zenodo.5437647" httpUri="https://zenodo.org/record/5437647/files/figure.png" pageId="35" pageNumber="482">Figs 19A</figureCitation>
,
<figureCitation id="132414B0EE15FFC18BC360DE3FA06E71" box="[1160,1210,1791,1813]" captionStart="Figure 21" captionStartId="74.[142,220,1755,1774]" captionTargetBox="[274,1290,197,1722]" captionTargetId="figure-153@74.[274,1291,197,1722]" captionTargetPageId="74" captionText="Figure 21. Style sac morphology; major typhlosole and ciliary tract (indicated by arrows). (A) Theodoxus fluviatilis. Scale bars (left to right) = 100 Mm, 10 Mm. (B) Neocyclotus dysoni ambiguum. Scale bars = 1 mm, 100 Mm. (C) Strombus mutabilis. Scale bars = 100 Mm, 10 Mm. (D) Crepidula plana. Scale bars = 100 Mm, 100 Mm. (E) Bithynia tentaculata. Scale bars = 1 mm, 100 Mm. (F) Cypraea robertsi. Scale bars = 1 mm, 100 Mm. (G) Conus jaspideus. Scale bars = 100 Mm, 10 Mm. (H) Ilyanassa obsoletus. Scale bars = 100 Mm, 10 Mm. Abbreviations: t1, major typhlosole; t2, minor typhlosole." figureDoi="http://doi.org/10.5281/zenodo.5437651" httpUri="https://zenodo.org/record/5437651/files/figure.png" pageId="35" pageNumber="482">21H</figureCitation>
). Typhlosoles flanking extensive region of transversely folded epithelium with differentiated cilia. Ciliary currents flowing clockwise within style sac. Crystalline style present.
</paragraph>
<paragraph id="8BA00835EE12FFC68FC566E53A726873" blockId="36.[142,759,196,279]" pageId="36" pageNumber="483">
<emphasis id="B96BD427EE12FFC68FC566E53BEF69BD" box="[142,245,196,217]" italics="true" pageId="36" pageNumber="483">Hindgut.</emphasis>
Intestine exiting style sac, curving dorsally and slightly posteriorly around rear wall of kidney. Anal gland absent.
</paragraph>
<paragraph id="8BA00835EE12FFC68FC5677C3A806817" blockId="36.[142,758,349,770]" box="[142,410,349,371]" pageId="36" pageNumber="483">
<emphasis id="B96BD427EE12FFC68FC5677C3A806817" box="[142,410,349,371]" italics="true" pageId="36" pageNumber="483">Reno-pericardial system</emphasis>
</paragraph>
<paragraph id="8BA00835EE12FFC68FC5675D3AAD6A65" blockId="36.[142,758,349,770]" pageId="36" pageNumber="483">
Kidney lumen elongate, projecting slightly into pallial cavity (
<figureCitation id="132414B0EE12FFC68FAE67BA3A5D68D5" box="[229,327,411,433]" captionStart="Figure 23" captionStartId="78.[142,220,1620,1639]" captionTargetBox="[323,1232,198,1588]" captionTargetId="figure-190@78.[310,1256,198,1616]" captionTargetPageId="78" captionText="Figure 23. Kidney morphology; derived caenogastropods with two differentiated lobes of lamellae. (A) Neverita duplicata, dorsal view. Kidney opened between lobes of differentiated lamellae. (B) Cypraea robertsi, dorsal view. Intestine removed to reveal internal details. (C) Panarona clavatula, dorsal view, posterior chamber on left, anterior chamber on right. Kidney roof opened between lobes of primary and secondary lamellae. Note meronephridial condition (no interdigitation of lamellae) and that dorsal branch of afferent renal vessel supplies both primary and secondary tubules. (D) Prunum apicinum, dorsal view. Note small region of secondary tubules within kidney roof supplied by dorsal branch of afferent renal vessel. (E) Ilyanassa obsoletus, dorsal view. Note pycnonephridial condition (interdigitation of lamellae). Scale bar = 1 mm. Abbreviations: al, anterior kidney lobe; arv, afferent renal vessel; au, auricle; darv, dorsal branch of afferent renal vessel; int, intestine; ng, nephridial gland; np, nephropore; per, pericardium; pl, posterior kidney lobe; rpc, reno-pericardial canal; ve, ventricle." figureDoi="http://doi.org/10.5281/zenodo.5437657" httpUri="https://zenodo.org/record/5437657/files/figure.png" pageId="36" pageNumber="483">Fig. 23E</figureCitation>
). Afferent renal vessel entering kidney floor anteriorly, to right of reno-pericardial canal (rpc).
<collectionCode id="ED0E90F0EE12FFC68F8567F93A096889" box="[206,275,472,493]" pageId="36" pageNumber="483">VARV</collectionCode>
extending posteriorly within floor, supplying numerous clusters of primary tubules along right wall.
<collectionCode id="ED0E90F0EE12FFC68F8764343A0F6B4E" box="[204,277,533,554]" pageId="36" pageNumber="483">DARV</collectionCode>
extending dorsally attached to front wall of kidney, turning posteriorly within centre of kidney roof, supplying secondary tubules. Primary and secondary tubules pycnonephridial, broadly interdigitating along right side of roof. Simple, narrow, thinwalled gland of Leiblein ampulla present within afferent renal vessel, extending within
<collectionCode id="ED0E90F0EE12FFC68D5164EC39796B86" box="[538,611,717,738]" pageId="36" pageNumber="483">DARV</collectionCode>
(
<figureCitation id="132414B0EE12FFC68D3964EC39DC6B87" box="[626,710,717,739]" captionStart="Figure 7" captionStartId="51.[155,233,1525,1544]" captionTargetBox="[373,1221,197,1490]" captionTargetId="figure-270@51.[363,1227,197,1493]" captionTargetPageId="51" captionText="Figure 7. Morphology of cephalic haemocoel among derived caenogastropods. (A) Panarona clavatula. Siphon obscured by left cephalic tentacle and is not visible in this view. Note large, modified left cephalic tentacle and large proboscis retractor connecting to columellar muscle. (B) Ilyanassa obsoletus. Note second visceral ganglion lying below pericardium and large radular retractors connecting to base of proboscis sheath; ampulla of gland of Leiblein visible within afferent renal vessel. Duct of gland of Leiblein stippled. (C) Nitidiscala tinctum. Note compartmentalized tissue supporting jaw and buccal ganglia lying between salivary glands where the latter enter buccal cavity. (D) Prunum apicinum. Note duct of poison gland opening mid-ventrally to base of buccal cavity, just behind opening of accessory salivary gland. (E) Conus jaspideus. Siphon not visible in this view. Note long coiled duct of poison gland; buccal ganglion visible where short arm of radular sac enters buccal cavity. Base of proboscis sheath is shown, but rhynchocoel has not been opened. Scale bar = 1 mm. Abbreviations: asg, accessory salivary gland; dasg, duct of accessory salivary gland; e, oesophagus; ec, oesophageal caecum; gl, gland of Leiblein; j, jaw; kd, kidney; nr, circum-oesophageal nerve ring; os, osphradium; per, pericardium; ps, proboscis sheath; rs, long arm of radular sac; sb, sub-oesophageal ganglion; sgd, salivary gland duct; sgl, salivary glands; si, siphon; sig, siphonal ganglion; sp, supra-oesophageal ganglion; tn, tentacular nerve; vg, visceral ganglion; vl, valve of Leiblein; z, zygoneury." figureDoi="http://doi.org/10.5281/zenodo.5437621" httpUri="https://zenodo.org/record/5437621/files/figure.png" pageId="36" pageNumber="483">Fig. 7B</figureCitation>
, gl). Nephridial gland present.
</paragraph>
<paragraph id="8BA00835EE12FFC68FC5656939276A39" blockId="36.[142,758,840,1445]" box="[142,573,840,861]" pageId="36" pageNumber="483">
<emphasis id="B96BD427EE12FFC68FC5656939276A39" box="[142,573,840,861]" italics="true" pageId="36" pageNumber="483">Nervous system and sensory structures</emphasis>
</paragraph>
<paragraph id="8BA00835EE12FFC68FC565463A306CC0" blockId="36.[142,758,840,1445]" pageId="36" pageNumber="483">
Nervous system epiathroid, right zygoneurous, left dialyneurous (
<figureCitation id="132414B0EE12FFC68E7265A43A876AFF" box="[313,413,901,923]" captionStart="Figure 24" captionStartId="81.[155,233,1702,1721]" captionTargetBox="[306,1287,200,1669]" captionTargetId="figure-148@81.[301,1290,192,1674]" captionTargetPageId="81" captionText="Figure 24. Circum-oesophageal nerve ring morphology. (A) Neocyclotus dysoni ambiguum. Scale bar = 0.5 mm. (B) Lampanella minima. Scale bar = 0.5 mm. (C) Crepidula plana. Scale bar = 0.5 mm. (D) Cypraea robertsi. Upper diagram depicts cerebral, pleural, supra-esophageal, and sub-oesophageal ganglia; lower diagram depicts pedal ganglia and pedal cords. Scale bar = 1 mm. (E) Nitidiscala tinctum. Scale bar = 0.25 mm. (F) Prunum apicinum. Scale bar = 0.5 mm. (G) Ilyanassa obsoletus. Scale bar = 0.5 mm. Abbreviations: bg, buccal ganglion; ceg, cerebral ganglion; d, dialyneury; e, oesophagus; pc, pedal cord; pdg, pedal ganglion; plg, pleural ganglion; ppg, propodial ganglion; sb, sub-oesophageal ganglion; sc, statocone; sp, supra-oesophageal ganglion; z, zygoneury." figureDoi="http://doi.org/10.5281/zenodo.5437659" httpUri="https://zenodo.org/record/5437659/files/figure.png" pageId="36" pageNumber="483">Fig. 24G</figureCitation>
). All ganglia except visceral ganglia concentrated in circum-oesophageal nerve ring. Cerebral (ceg) and pleural (plg) ganglia broadly fused. Single nerve splitting near nerve ring into tentacular and optic nerves. Tentacular nerve giving off several small branches before splitting into two nerves of equal size, extending length of tentacle (
<figureCitation id="132414B0EE12FFC68D3B621C39D96D37" box="[624,707,1085,1107]" captionStart="Figure 7" captionStartId="51.[155,233,1525,1544]" captionTargetBox="[373,1221,197,1490]" captionTargetId="figure-270@51.[363,1227,197,1493]" captionTargetPageId="51" captionText="Figure 7. Morphology of cephalic haemocoel among derived caenogastropods. (A) Panarona clavatula. Siphon obscured by left cephalic tentacle and is not visible in this view. Note large, modified left cephalic tentacle and large proboscis retractor connecting to columellar muscle. (B) Ilyanassa obsoletus. Note second visceral ganglion lying below pericardium and large radular retractors connecting to base of proboscis sheath; ampulla of gland of Leiblein visible within afferent renal vessel. Duct of gland of Leiblein stippled. (C) Nitidiscala tinctum. Note compartmentalized tissue supporting jaw and buccal ganglia lying between salivary glands where the latter enter buccal cavity. (D) Prunum apicinum. Note duct of poison gland opening mid-ventrally to base of buccal cavity, just behind opening of accessory salivary gland. (E) Conus jaspideus. Siphon not visible in this view. Note long coiled duct of poison gland; buccal ganglion visible where short arm of radular sac enters buccal cavity. Base of proboscis sheath is shown, but rhynchocoel has not been opened. Scale bar = 1 mm. Abbreviations: asg, accessory salivary gland; dasg, duct of accessory salivary gland; e, oesophagus; ec, oesophageal caecum; gl, gland of Leiblein; j, jaw; kd, kidney; nr, circum-oesophageal nerve ring; os, osphradium; per, pericardium; ps, proboscis sheath; rs, long arm of radular sac; sb, sub-oesophageal ganglion; sgd, salivary gland duct; sgl, salivary glands; si, siphon; sig, siphonal ganglion; sp, supra-oesophageal ganglion; tn, tentacular nerve; vg, visceral ganglion; vl, valve of Leiblein; z, zygoneury." figureDoi="http://doi.org/10.5281/zenodo.5437621" httpUri="https://zenodo.org/record/5437621/files/figure.png" pageId="36" pageNumber="483">Fig. 7B</figureCitation>
, tn). Buccal ganglia (
<figureCitation id="132414B0EE12FFC68E0D627D3AB26D15" box="[326,424,1116,1138]" captionStart="Figure 24" captionStartId="81.[155,233,1702,1721]" captionTargetBox="[306,1287,200,1669]" captionTargetId="figure-148@81.[301,1290,192,1674]" captionTargetPageId="81" captionText="Figure 24. Circum-oesophageal nerve ring morphology. (A) Neocyclotus dysoni ambiguum. Scale bar = 0.5 mm. (B) Lampanella minima. Scale bar = 0.5 mm. (C) Crepidula plana. Scale bar = 0.5 mm. (D) Cypraea robertsi. Upper diagram depicts cerebral, pleural, supra-esophageal, and sub-oesophageal ganglia; lower diagram depicts pedal ganglia and pedal cords. Scale bar = 1 mm. (E) Nitidiscala tinctum. Scale bar = 0.25 mm. (F) Prunum apicinum. Scale bar = 0.5 mm. (G) Ilyanassa obsoletus. Scale bar = 0.5 mm. Abbreviations: bg, buccal ganglion; ceg, cerebral ganglion; d, dialyneury; e, oesophagus; pc, pedal cord; pdg, pedal ganglion; plg, pleural ganglion; ppg, propodial ganglion; sb, sub-oesophageal ganglion; sc, statocone; sp, supra-oesophageal ganglion; z, zygoneury." figureDoi="http://doi.org/10.5281/zenodo.5437659" httpUri="https://zenodo.org/record/5437659/files/figure.png" pageId="36" pageNumber="483">Fig. 24G</figureCitation>
, bg) present at anterior ends of cerebral ganglia. Large siphonal ganglion present (
<figureCitation id="132414B0EE12FFC68FDD62B83BF36DCB" box="[150,233,1177,1199]" captionStart="Figure 7" captionStartId="51.[155,233,1525,1544]" captionTargetBox="[373,1221,197,1490]" captionTargetId="figure-270@51.[363,1227,197,1493]" captionTargetPageId="51" captionText="Figure 7. Morphology of cephalic haemocoel among derived caenogastropods. (A) Panarona clavatula. Siphon obscured by left cephalic tentacle and is not visible in this view. Note large, modified left cephalic tentacle and large proboscis retractor connecting to columellar muscle. (B) Ilyanassa obsoletus. Note second visceral ganglion lying below pericardium and large radular retractors connecting to base of proboscis sheath; ampulla of gland of Leiblein visible within afferent renal vessel. Duct of gland of Leiblein stippled. (C) Nitidiscala tinctum. Note compartmentalized tissue supporting jaw and buccal ganglia lying between salivary glands where the latter enter buccal cavity. (D) Prunum apicinum. Note duct of poison gland opening mid-ventrally to base of buccal cavity, just behind opening of accessory salivary gland. (E) Conus jaspideus. Siphon not visible in this view. Note long coiled duct of poison gland; buccal ganglion visible where short arm of radular sac enters buccal cavity. Base of proboscis sheath is shown, but rhynchocoel has not been opened. Scale bar = 1 mm. Abbreviations: asg, accessory salivary gland; dasg, duct of accessory salivary gland; e, oesophagus; ec, oesophageal caecum; gl, gland of Leiblein; j, jaw; kd, kidney; nr, circum-oesophageal nerve ring; os, osphradium; per, pericardium; ps, proboscis sheath; rs, long arm of radular sac; sb, sub-oesophageal ganglion; sgd, salivary gland duct; sgl, salivary glands; si, siphon; sig, siphonal ganglion; sp, supra-oesophageal ganglion; tn, tentacular nerve; vg, visceral ganglion; vl, valve of Leiblein; z, zygoneury." figureDoi="http://doi.org/10.5281/zenodo.5437621" httpUri="https://zenodo.org/record/5437621/files/figure.png" pageId="36" pageNumber="483">Fig. 7B</figureCitation>
, sig) at dialyneury between nerve from left pleural and branch from anterior osphradial nerve. Posterior osphradial nerve giving rise to left arm of visceral loop. Propodial ganglia (
<figureCitation id="132414B0EE12FFC68D0E62D439B26C6E" box="[581,680,1269,1291]" captionStart="Figure 24" captionStartId="81.[155,233,1702,1721]" captionTargetBox="[306,1287,200,1669]" captionTargetId="figure-148@81.[301,1290,192,1674]" captionTargetPageId="81" captionText="Figure 24. Circum-oesophageal nerve ring morphology. (A) Neocyclotus dysoni ambiguum. Scale bar = 0.5 mm. (B) Lampanella minima. Scale bar = 0.5 mm. (C) Crepidula plana. Scale bar = 0.5 mm. (D) Cypraea robertsi. Upper diagram depicts cerebral, pleural, supra-esophageal, and sub-oesophageal ganglia; lower diagram depicts pedal ganglia and pedal cords. Scale bar = 1 mm. (E) Nitidiscala tinctum. Scale bar = 0.25 mm. (F) Prunum apicinum. Scale bar = 0.5 mm. (G) Ilyanassa obsoletus. Scale bar = 0.5 mm. Abbreviations: bg, buccal ganglion; ceg, cerebral ganglion; d, dialyneury; e, oesophagus; pc, pedal cord; pdg, pedal ganglion; plg, pleural ganglion; ppg, propodial ganglion; sb, sub-oesophageal ganglion; sc, statocone; sp, supra-oesophageal ganglion; z, zygoneury." figureDoi="http://doi.org/10.5281/zenodo.5437659" httpUri="https://zenodo.org/record/5437659/files/figure.png" pageId="36" pageNumber="483">Fig. 24G</figureCitation>
, ppg) present at anterior ends of pedal ganglia (pdg). Paired, asymmetrical visceral ganglia (
<figureCitation id="132414B0EE12FFC68EBE631239536C2C" box="[501,585,1331,1353]" captionStart="Figure 7" captionStartId="51.[155,233,1525,1544]" captionTargetBox="[373,1221,197,1490]" captionTargetId="figure-270@51.[363,1227,197,1493]" captionTargetPageId="51" captionText="Figure 7. Morphology of cephalic haemocoel among derived caenogastropods. (A) Panarona clavatula. Siphon obscured by left cephalic tentacle and is not visible in this view. Note large, modified left cephalic tentacle and large proboscis retractor connecting to columellar muscle. (B) Ilyanassa obsoletus. Note second visceral ganglion lying below pericardium and large radular retractors connecting to base of proboscis sheath; ampulla of gland of Leiblein visible within afferent renal vessel. Duct of gland of Leiblein stippled. (C) Nitidiscala tinctum. Note compartmentalized tissue supporting jaw and buccal ganglia lying between salivary glands where the latter enter buccal cavity. (D) Prunum apicinum. Note duct of poison gland opening mid-ventrally to base of buccal cavity, just behind opening of accessory salivary gland. (E) Conus jaspideus. Siphon not visible in this view. Note long coiled duct of poison gland; buccal ganglion visible where short arm of radular sac enters buccal cavity. Base of proboscis sheath is shown, but rhynchocoel has not been opened. Scale bar = 1 mm. Abbreviations: asg, accessory salivary gland; dasg, duct of accessory salivary gland; e, oesophagus; ec, oesophageal caecum; gl, gland of Leiblein; j, jaw; kd, kidney; nr, circum-oesophageal nerve ring; os, osphradium; per, pericardium; ps, proboscis sheath; rs, long arm of radular sac; sb, sub-oesophageal ganglion; sgd, salivary gland duct; sgl, salivary glands; si, siphon; sig, siphonal ganglion; sp, supra-oesophageal ganglion; tn, tentacular nerve; vg, visceral ganglion; vl, valve of Leiblein; z, zygoneury." figureDoi="http://doi.org/10.5281/zenodo.5437621" httpUri="https://zenodo.org/record/5437621/files/figure.png" pageId="36" pageNumber="483">Fig. 7B</figureCitation>
, vg) straddling posterior oesophagus. Highly asymmetrical statocysts with single statoliths present ventral to nerve ring (
<figureCitation id="132414B0EE12FFC68FDD63AE3BE26CC0" box="[150,248,1423,1445]" captionStart="Figure 24" captionStartId="81.[155,233,1702,1721]" captionTargetBox="[306,1287,200,1669]" captionTargetId="figure-148@81.[301,1290,192,1674]" captionTargetPageId="81" captionText="Figure 24. Circum-oesophageal nerve ring morphology. (A) Neocyclotus dysoni ambiguum. Scale bar = 0.5 mm. (B) Lampanella minima. Scale bar = 0.5 mm. (C) Crepidula plana. Scale bar = 0.5 mm. (D) Cypraea robertsi. Upper diagram depicts cerebral, pleural, supra-esophageal, and sub-oesophageal ganglia; lower diagram depicts pedal ganglia and pedal cords. Scale bar = 1 mm. (E) Nitidiscala tinctum. Scale bar = 0.25 mm. (F) Prunum apicinum. Scale bar = 0.5 mm. (G) Ilyanassa obsoletus. Scale bar = 0.5 mm. Abbreviations: bg, buccal ganglion; ceg, cerebral ganglion; d, dialyneury; e, oesophagus; pc, pedal cord; pdg, pedal ganglion; plg, pleural ganglion; ppg, propodial ganglion; sb, sub-oesophageal ganglion; sc, statocone; sp, supra-oesophageal ganglion; z, zygoneury." figureDoi="http://doi.org/10.5281/zenodo.5437659" httpUri="https://zenodo.org/record/5437659/files/figure.png" pageId="36" pageNumber="483">Fig. 24G</figureCitation>
, sc).
</paragraph>
<paragraph id="8BA00835EE12FFC68FC563CB3BEA6C9B" blockId="36.[142,758,1514,1905]" box="[142,240,1514,1535]" pageId="36" pageNumber="483">
<emphasis id="B96BD427EE12FFC68FC563CB3BEA6C9B" box="[142,240,1514,1535]" italics="true" pageId="36" pageNumber="483">Remarks</emphasis>
</paragraph>
<paragraph id="8BA00835EE12FFC68FC5602838C66852" blockId="36.[142,758,1514,1905]" lastBlockId="36.[806,1422,196,555]" pageId="36" pageNumber="483">
<bibRefCitation id="EF8E75C4EE12FFC68FC560283A326F7B" author="Dimon AC" box="[142,296,1545,1567]" pageId="36" pageNumber="483" pagination="1 - 48" refId="ref73556" refString="Dimon AC. 1905. The mud snail: Nassa obsoleta. Cold Spring Harbor Monographs 5: 1 - 48." type="journal article" year="1905">Dimon (1905)</bibRefCitation>
provided some preliminary observations on the external anatomy and mantle cavity, reproductive and alimentary systems.
<bibRefCitation id="EF8E75C4EE12FFC68EB2606639836F39" author="Brown SC" box="[505,665,1607,1629]" pageId="36" pageNumber="483" pagination="447 - 500" refId="ref72751" refString="Brown SC. 1969. The structure and function of the digestive system of the mud snail Nassarius obsoletus (Say). Malacologia 9: 447 - 500." type="journal article" year="1969">Brown (1969)</bibRefCitation>
gave a more detailed description of the alimentary system. The present description is congruent with that of Brown in many respects with several minor exceptions. For example, Brown described the salivary gland ducts as entering the buccal cavity dorso-laterally. Histological sections revealed here that fine ducts, embedded within the walls of the buccal mass, enter ventrally under the odontophore. Brown figured the gland of Leiblein as lacking a terminal ampulla. The ampulla was found here to enter the dorsal branch of the afferent renal vessel and is thin-walled and simpler than those in some muricids (e.g.
<bibRefCitation id="EF8E75C4EE12FFC68C6D670138D56852" author="Carriker MR" box="[806,975,288,310]" pageId="36" pageNumber="483" pagination="441 - 506" refId="ref72904" refString="Carriker MR. 1943. On the structure and function of the proboscis in the common oyster drill, Urosalpinx cinerea Say. Journal of Morphology 73: 441 - 506." type="journal article" year="1943">Carriker, 1943</bibRefCitation>
).
</paragraph>
<paragraph id="8BA00835EE12FFC68C74671E38D26B4F" blockId="36.[806,1422,196,555]" pageId="36" pageNumber="483">
Within the midgut,
<bibRefCitation id="EF8E75C4EE12FFC68B6A671E3FA16831" author="Brown SC" box="[1057,1211,319,341]" pageId="36" pageNumber="483" pagination="447 - 500" refId="ref72751" refString="Brown SC. 1969. The structure and function of the digestive system of the mud snail Nassarius obsoletus (Say). Malacologia 9: 447 - 500." type="journal article" year="1969">Brown (1969)</bibRefCitation>
indicated that ciliary currents adjacent to the intestinal groove flow towards the intestine. Here, these currents were found to flow transversely in the direction of the sorting area or, more posteriorly, toward the oesophageal aperture itself. In addition, Brown did not acknowledge the presence of an elongate glandular fold bounding the gastric shield.
</paragraph>
<paragraph id="8BA00835EE12FFC68C6D645338876BE3" blockId="36.[806,1422,626,1905]" box="[806,925,626,647]" pageId="36" pageNumber="483">
<emphasis id="B96BD427EE12FFC68C6D645338876BE3" box="[806,925,626,647]" italics="true" pageId="36" pageNumber="483">Discussion</emphasis>
</paragraph>
<paragraph id="8BA00835EE12FFC68C6D64B13E226A5B" blockId="36.[806,1422,626,1905]" pageId="36" pageNumber="483">
Like other nassariids,
<taxonomicName id="4C1F73B6EE12FFC68B6C64B13E1E6BC1" baseAuthorityName="SAY" baseAuthorityYear="1822" box="[1063,1284,656,677]" class="Gastropoda" family="Nassariidae" genus="Ilyanassa" higherTaxonomySource="GBIF" kingdom="Animalia" order="Neogastropoda" pageId="36" pageNumber="483" phylum="Mollusca" rank="species" species="obsoletus">
<emphasis id="B96BD427EE12FFC68B6C64B13E1E6BC1" box="[1063,1284,656,677]" italics="true" pageId="36" pageNumber="483">Ilyanassa obsoletus</emphasis>
</taxonomicName>
bears weak dorsal folds in the anterior oesophagus (= glande fromboisée) (
<bibRefCitation id="EF8E75C4EE12FFC68CC164EC3F2C6B87" author="Graham A" box="[906,1078,717,739]" pageId="36" pageNumber="483" pagination="1 - 23" refId="ref74286" refString="Graham A. 1941. The oesophagus of the stenoglossan prosobranchs. Proceedings of the Royal Society of Edinburgh 61: 1 - 23." type="journal article" year="1941">Graham, 1941</bibRefCitation>
). A valve of Leiblein and a small gland of Leiblein are present; accessory salivary glands and the anal gland are lacking (
<bibRefCitation id="EF8E75C4EE12FFC68B96652A3E986A45" author="Graham A" box="[1245,1410,779,801]" pageId="36" pageNumber="483" pagination="1 - 23" refId="ref74286" refString="Graham A. 1941. The oesophagus of the stenoglossan prosobranchs. Proceedings of the Royal Society of Edinburgh 61: 1 - 23." type="journal article" year="1941">Graham, 1941</bibRefCitation>
). The kidney is pycnonephridial (
<bibRefCitation id="EF8E75C4EE12FFC68BD865083E316A5B" author="Ponder WF" box="[1171,1323,809,831]" pageId="36" pageNumber="483" pagination="295 - 338" refId="ref78484" refString="Ponder WF. 1973. The origin and evolution of the Neogastropoda. Malacologia 12: 295 - 338." type="journal article" year="1973">Ponder, 1973</bibRefCitation>
).
</paragraph>
<paragraph id="8BA00835EE12FFC68C7465693FCF6C4E" blockId="36.[806,1422,626,1905]" pageId="36" pageNumber="483">
Female reproductive anatomy of
<taxonomicName id="4C1F73B6EE12FFC68BF965693E976A39" baseAuthorityName="SAY" baseAuthorityYear="1822" box="[1202,1421,840,861]" class="Gastropoda" family="Nassariidae" genus="Ilyanassa" higherTaxonomySource="GBIF" kingdom="Animalia" order="Neogastropoda" pageId="36" pageNumber="483" phylum="Mollusca" rank="species" species="obsoletus">
<emphasis id="B96BD427EE12FFC68BF965693E976A39" box="[1202,1421,840,861]" italics="true" pageId="36" pageNumber="483">Ilyanassa obsoletus</emphasis>
</taxonomicName>
is consistent with described nassariids in the presence of a gonopericardial connection and an ingesting gland, but is unique in lacking a bursa (
<bibRefCitation id="EF8E75C4EE12FFC68BB965853E936ADD" author="Fretter V" box="[1266,1417,932,953]" pageId="36" pageNumber="483" pagination="173 - 211" refId="ref73747" refString="Fretter V. 1941. The genital ducts of some British stenoglossan prosobranchs. Journal of the Marine Biological Association (United Kingdom) 25: 173 - 211." type="journal article" year="1941">Fretter, 1941</bibRefCitation>
;
<bibRefCitation id="EF8E75C4EE12FFC68C6D65E238F56ABC" author="Johansson J" box="[806,1007,962,985]" pageId="36" pageNumber="483" pagination="81 - 91" refId="ref76171" refString="Johansson J. 1957. Notes on the littorinacean and stenoglossan genital organs, and a comparison with the Rissoacea. Zoologiska Bidrag fran Uppsala 32: 81 - 91." type="journal article" year="1957">Johansson, 1957</bibRefCitation>
;
<bibRefCitation id="EF8E75C4EE12FFC68B4F65E23FAD6ABD" author="Houston RS" box="[1028,1207,963,985]" pageId="36" pageNumber="483" pagination="27 - 46" refId="ref75881" refString="Houston RS. 1976. The structure and function of neogastropod reproductive systems: with special reference to Columbella fuscata Sowerby, 1832. The Veliger 19: 27 - 46." type="journal article" year="1976">Houston, 1976</bibRefCitation>
).
<bibRefCitation id="EF8E75C4EE12FFC68B9F65E23E946ABD" author="Houston RS" box="[1236,1422,963,985]" pageId="36" pageNumber="483" pagination="27 - 46" refId="ref75881" refString="Houston RS. 1976. The structure and function of neogastropod reproductive systems: with special reference to Columbella fuscata Sowerby, 1832. The Veliger 19: 27 - 46." type="journal article" year="1976">Houston (1976)</bibRefCitation>
described the presence of an opening within the oviduct to the mantle cavity in
<taxonomicName id="4C1F73B6EE12FFC68B2C62203E7E6D72" baseAuthorityName="Smith" baseAuthorityYear="1967" box="[1127,1380,1025,1046]" class="Gastropoda" family="Nassariidae" genus="Nassarius" kingdom="Animalia" order="Neogastropoda" pageId="36" pageNumber="483" phylum="Mollusca" rank="species" species="incrassatus">
<emphasis id="B96BD427EE12FFC68B2C62203E7E6D72" box="[1127,1380,1025,1046]" italics="true" pageId="36" pageNumber="483">Nassarius incrassatus</emphasis>
</taxonomicName>
, an aperture recorded only in several muricids (
<bibRefCitation id="EF8E75C4EE12FFC68A6F623E38446D37" author="Houston RS" pageId="36" pageNumber="483" pagination="27 - 46" refId="ref75881" refString="Houston RS. 1976. The structure and function of neogastropod reproductive systems: with special reference to Columbella fuscata Sowerby, 1832. The Veliger 19: 27 - 46." type="journal article" year="1976">Houston, 1976</bibRefCitation>
) and unknown among nassariids.
<bibRefCitation id="EF8E75C4EE12FFC68BAB621F3E946D37" author="Houston RS" box="[1248,1422,1085,1107]" pageId="36" pageNumber="483" pagination="27 - 46" refId="ref75881" refString="Houston RS. 1976. The structure and function of neogastropod reproductive systems: with special reference to Columbella fuscata Sowerby, 1832. The Veliger 19: 27 - 46." type="journal article" year="1976">Houston (1976)</bibRefCitation>
did not find a seminal receptacle or an ingesting gland, structures reported as present in a different analysis of the same species (
<bibRefCitation id="EF8E75C4EE12FFC68B6F62B83FF36DCB" author="Johansson J" box="[1060,1257,1177,1199]" pageId="36" pageNumber="483" pagination="81 - 91" refId="ref76171" refString="Johansson J. 1957. Notes on the littorinacean and stenoglossan genital organs, and a comparison with the Rissoacea. Zoologiska Bidrag fran Uppsala 32: 81 - 91." type="journal article" year="1957">Johansson, 1957</bibRefCitation>
). The present study noted several outpocketings of the albumen gland. Although their function could not be confirmed, similar outpocketings are known to store sperm in other neogastropods (e.g.
<bibRefCitation id="EF8E75C4EE12FFC68B0763353FD36C4E" author="Kool SP" box="[1100,1225,1300,1322]" pageId="36" pageNumber="483" pagination="155 - 259" refId="ref76556" refString="Kool SP. 1993. Phylogenetic analysis of the Rapaninae (Neogastropoda: Muricidae). Malacologia 35: 155 - 259." type="journal article" year="1993">Kool, 1993</bibRefCitation>
).
</paragraph>
<paragraph id="8BA00835EE12FFC68C74631238766F65" blockId="36.[806,1422,626,1905]" pageId="36" pageNumber="483">
In male nassariids, a diverticulum connecting the base of the prostate with the mantle lumen may be open (
<bibRefCitation id="EF8E75C4EE12FFC68C3A63513F176CE2" author="Fretter V" box="[881,1037,1392,1414]" pageId="36" pageNumber="483" pagination="173 - 211" refId="ref73747" refString="Fretter V. 1941. The genital ducts of some British stenoglossan prosobranchs. Journal of the Marine Biological Association (United Kingdom) 25: 173 - 211." type="journal article" year="1941">Fretter, 1941</bibRefCitation>
), fused shut (present study), or lacking entirely (
<bibRefCitation id="EF8E75C4EE12FFC68CB063AE3FB76CC1" author="Houston RS" box="[1019,1197,1423,1445]" pageId="36" pageNumber="483" pagination="27 - 46" refId="ref75881" refString="Houston RS. 1976. The structure and function of neogastropod reproductive systems: with special reference to Columbella fuscata Sowerby, 1832. The Veliger 19: 27 - 46." type="journal article" year="1976">Houston, 1976</bibRefCitation>
). The penial duct within the dorso-ventrally flattened penis may be subcentral (present study) or centrally located (
<bibRefCitation id="EF8E75C4EE12FFC68A7D63ED387A6F65" author="Fretter V" pageId="36" pageNumber="483" pagination="173 - 211" refId="ref73747" refString="Fretter V. 1941. The genital ducts of some British stenoglossan prosobranchs. Journal of the Marine Biological Association (United Kingdom) 25: 173 - 211." type="journal article" year="1941">Fretter, 1941</bibRefCitation>
).
</paragraph>
<paragraph id="8BA00835EE12FFC78C7460283AF36B2E" blockId="36.[806,1422,626,1905]" lastBlockId="37.[155,771,196,954]" lastPageId="37" lastPageNumber="484" pageId="36" pageNumber="483">
There appear to be
<specimenCount id="9D19C3BCEE12FFC68B52602B3F9F6F7B" box="[1049,1157,1546,1567]" pageId="36" pageNumber="483" type="generic" typeStatus="types">two types</specimenCount>
of midgut organization within the
<taxonomicName id="4C1F73B6EE12FFC68CF160093F5C6F5A" authorityName="Iredale" authorityYear="1916" box="[954,1094,1576,1598]" class="Gastropoda" family="Nassariidae" kingdom="Animalia" order="Neogastropoda" pageId="36" pageNumber="483" phylum="Mollusca" rank="family">Nassariidae</taxonomicName>
. The midgut structure of
<taxonomicName id="4C1F73B6EE12FFC68C6D60663F646F39" authority="(Smith, 1967)" baseAuthorityName="Smith" baseAuthorityYear="1967" box="[806,1150,1607,1629]" class="Gastropoda" family="Nassariidae" genus="Nassarius" kingdom="Animalia" order="Neogastropoda" pageId="36" pageNumber="483" phylum="Mollusca" rank="species" species="incrassatus">
<emphasis id="B96BD427EE12FFC68C6D606638DD6F38" box="[806,967,1607,1628]" italics="true" pageId="36" pageNumber="483">N. incrassatus</emphasis>
(
<bibRefCitation id="EF8E75C4EE12FFC68CAA60663F6C6F39" author="Smith EH" box="[993,1142,1607,1629]" pageId="36" pageNumber="483" pagination="23 - 42" refId="ref80043" refString="Smith EH. 1967. The neogastropod midgut, with notes on the digestive diverticula and intestine. Transactions of the Royal Society of Edinburgh 67: 23 - 42." type="journal article" year="1967">Smith, 1967</bibRefCitation>
)
</taxonomicName>
and
<taxonomicName id="4C1F73B6EE12FFC68B8160663F176F1F" authority="(Medinskaya, 1993)" baseAuthorityName="Medinskaya" baseAuthorityYear="1993" class="Gastropoda" family="Nassariidae" genus="Tritia" kingdom="Animalia" order="Neogastropoda" pageId="36" pageNumber="483" phylum="Mollusca" rank="species" species="fratercula">
<emphasis id="B96BD427EE12FFC68B8160663E976F38" box="[1226,1421,1607,1628]" italics="true" pageId="36" pageNumber="483">Tritia fratercula</emphasis>
(
<bibRefCitation id="EF8E75C4EE12FFC68C6660443F1F6F1F" author="Medinskaya AI" box="[813,1029,1637,1659]" pageId="36" pageNumber="483" pagination="17 - 24" refId="ref77339" refString="Medinskaya AI. 1993. Anatomy of the midgut of some Neogastropoda from the offshore zone of the Japan Sea. Ruthenica 3: 17 - 24." type="journal article" year="1993">Medinskaya, 1993</bibRefCitation>
)
</taxonomicName>
is congruent in many respects with that of
<taxonomicName id="4C1F73B6EE12FFC68CF560A53F876FFD" baseAuthorityName="SAY" baseAuthorityYear="1822" box="[958,1181,1668,1689]" class="Gastropoda" family="Nassariidae" genus="Ilyanassa" higherTaxonomySource="GBIF" kingdom="Animalia" order="Neogastropoda" pageId="36" pageNumber="483" phylum="Mollusca" rank="species" species="obsoletus">
<emphasis id="B96BD427EE12FFC68CF560A53F876FFD" box="[958,1181,1668,1689]" italics="true" pageId="36" pageNumber="483">Ilyanassa obsoletus</emphasis>
</taxonomicName>
, with several minor exceptions. For example, in the two former taxa, the depression bearing the paired ducts of the digestive gland is more pronounced than in
<taxonomicName id="4C1F73B6EE12FFC68BE760C03E306F91" baseAuthorityName="SAY" baseAuthorityYear="1822" box="[1196,1322,1760,1782]" class="Gastropoda" family="Nassariidae" genus="Ilyanassa" higherTaxonomySource="GBIF" kingdom="Animalia" order="Neogastropoda" pageId="36" pageNumber="483" phylum="Mollusca" rank="species" species="obsoletus">
<emphasis id="B96BD427EE12FFC68BE760C03E306F91" box="[1196,1322,1760,1782]" italics="true" pageId="36" pageNumber="483">I. obsoletus</emphasis>
</taxonomicName>
, and the sorting area to the left of the oesophagus is lacking. The ‘ciliary sorting area’ (Smith) of
<taxonomicName id="4C1F73B6EE12FFC68B84613F3E6A6E57" baseAuthorityName="Smith" baseAuthorityYear="1967" box="[1231,1392,1822,1843]" class="Gastropoda" family="Nassariidae" genus="Nassarius" kingdom="Animalia" order="Neogastropoda" pageId="36" pageNumber="483" phylum="Mollusca" rank="species" species="incrassatus">
<emphasis id="B96BD427EE12FFC68B84613F3E6A6E57" box="[1231,1392,1822,1843]" italics="true" pageId="36" pageNumber="483">N. incrassatus</emphasis>
</taxonomicName>
is homologous to the style sac region, and the ‘posterior sorting area’, occurring ventrally in front of the intestinal groove, has no equivalent in
<taxonomicName id="4C1F73B6EE13FFC78D0D66E439D969BD" baseAuthorityName="SAY" baseAuthorityYear="1822" box="[582,707,196,218]" class="Gastropoda" family="Nassariidae" genus="Ilyanassa" higherTaxonomySource="GBIF" kingdom="Animalia" order="Neogastropoda" pageId="37" pageNumber="484" phylum="Mollusca" rank="species" species="obsoletus">
<emphasis id="B96BD427EE13FFC78D0D66E439D969BD" box="[582,707,196,218]" italics="true" pageId="37" pageNumber="484">I. obsoletus</emphasis>
</taxonomicName>
. The ‘posterior sorting area’ (Medinskaya) of
<taxonomicName id="4C1F73B6EE13FFC78D1666C239F3699C" baseAuthorityName="Medinskaya" baseAuthorityYear="1993" box="[605,745,227,248]" class="Gastropoda" family="Nassariidae" genus="Tritia" kingdom="Animalia" order="Neogastropoda" pageId="37" pageNumber="484" phylum="Mollusca" rank="species" species="fratercula">
<emphasis id="B96BD427EE13FFC78D1666C239F3699C" box="[605,745,227,248]" italics="true" pageId="37" pageNumber="484">T. fratercula</emphasis>
</taxonomicName>
is not equivalent to the structure of the same name in
<taxonomicName id="4C1F73B6EE13FFC78FD067003A266852" baseAuthorityName="Smith" baseAuthorityYear="1967" box="[155,316,289,310]" class="Gastropoda" family="Nassariidae" genus="Nassarius" kingdom="Animalia" order="Neogastropoda" pageId="37" pageNumber="484" phylum="Mollusca" rank="species" species="incrassatus">
<emphasis id="B96BD427EE13FFC78FD067003A266852" box="[155,316,289,310]" italics="true" pageId="37" pageNumber="484">N. incrassatus</emphasis>
</taxonomicName>
, but is homologous to the style sac region. In contrast, the midgut organization of of
<taxonomicName id="4C1F73B6EE13FFC78D86671E39076817" authority="(Graham, 1949)" baseAuthorityName="Graham" baseAuthorityYear="1949" class="Gastropoda" family="Nassariidae" genus="Nassarius" kingdom="Animalia" order="Neogastropoda" pageId="37" pageNumber="484" phylum="Mollusca" rank="species" species="reticulatus">
<emphasis id="B96BD427EE13FFC78D86671E3A786816" italics="true" pageId="37" pageNumber="484">Nassarius reticulatus</emphasis>
(
<bibRefCitation id="EF8E75C4EE13FFC78E3A677C390F6817" author="Graham A" box="[369,533,349,371]" pageId="37" pageNumber="484" pagination="737 - 778" refId="ref74311" refString="Graham A. 1949. The molluscan midgut. Transactions of the Royal Society of Edinburgh 61: 737 - 778." type="journal article" year="1949">Graham, 1949</bibRefCitation>
)
</taxonomicName>
and
<taxonomicName id="4C1F73B6EE13FFC78D1E677C3A5268F6" authority="(Morton, 1960)" baseAuthorityName="Morton" baseAuthorityYear="1960" class="Gastropoda" family="Nassariidae" genus="Cyclope" higherTaxonomySource="GBIF" kingdom="Animalia" order="Neogastropoda" pageId="37" pageNumber="484" phylum="Mollusca" rank="species" species="neritea">
<emphasis id="B96BD427EE13FFC78D1E677C38186817" box="[597,770,349,371]" italics="true" pageId="37" pageNumber="484">Cyclope neritea</emphasis>
(
<bibRefCitation id="EF8E75C4EE13FFC78FE9675D3A5B68F6" author="Morton JE" box="[162,321,380,402]" pageId="37" pageNumber="484" pagination="96 - 105" refId="ref77744" refString="Morton JE. 1960. The habits of Cyclope neritea, a style-bearing stenoglossan gastropod. Proceedings of the Malacological Society of London 34: 96 - 105." type="journal article" year="1960">Morton, 1960</bibRefCitation>
)
</taxonomicName>
is characterized by an elongate fold bounding the oesophageal aperture to the left, separating the oesophagus from the sorting area; this fold is lacking in the other nassariids. In addition, the digestive gland ducts open to the sorting area to the left of the oesophagus and there is no fold bounding the gastric shield on the left.
</paragraph>
<paragraph id="8BA00835EE13FFC78FF8647239A06ADE" blockId="37.[155,771,196,954]" pageId="37" pageNumber="484">
The nervous system of
<taxonomicName id="4C1F73B6EE13FFC78E86647239B76B0C" baseAuthorityName="SAY" baseAuthorityYear="1822" box="[461,685,595,616]" class="Gastropoda" family="Nassariidae" genus="Ilyanassa" higherTaxonomySource="GBIF" kingdom="Animalia" order="Neogastropoda" pageId="37" pageNumber="484" phylum="Mollusca" rank="species" species="obsoletus">
<emphasis id="B96BD427EE13FFC78E86647239B76B0C" box="[461,685,595,616]" italics="true" pageId="37" pageNumber="484">Ilyanassa obsoletus</emphasis>
</taxonomicName>
differs from described nassariids only in minor detail (
<bibRefCitation id="EF8E75C4EE13FFC78DFB64503BCE6BC2" author="Risbec J" pageId="37" pageNumber="484" pagination="487 - 495" refId="ref79193" refString="Risbec J. 1952. Observations sur l'anatomie des Nassidae de Nouvelle-Caledonie. Bulletin de la Societe Zoologique de France 79: 487 - 495." type="journal article" year="1952">Risbec, 1952</bibRefCitation>
;
<bibRefCitation id="EF8E75C4EE13FFC78FAD64B1390C6BC2" author="Fretter V &amp; Graham A" box="[230,534,656,678]" pageId="37" pageNumber="484" refId="ref73894" refString="Fretter V, Graham A. 1962. British prosobranch molluscs; their functional anatomy and ecology. London: Ray Society." type="book" year="1962">Fretter &amp; Graham, 1962</bibRefCitation>
;
<bibRefCitation id="EF8E75C4EE13FFC78D6364B139D86BC2" author="Brown AC" box="[552,706,656,678]" pageId="37" pageNumber="484" pagination="309 - 361" refId="ref72781" refString="Brown AC. 1982. The biology of sandy-beach whelks of the genus Bullia (Nassariidae). Oceanography and Marine Biology: an Annual Review 20: 309 - 361." type="journal article" year="1982">Brown, 1982</bibRefCitation>
). For example, in
<taxonomicName id="4C1F73B6EE13FFC78E64648E3AB76BA0" baseAuthorityName="SAY" baseAuthorityYear="1822" box="[303,429,687,708]" class="Gastropoda" family="Nassariidae" genus="Ilyanassa" higherTaxonomySource="GBIF" kingdom="Animalia" order="Neogastropoda" pageId="37" pageNumber="484" phylum="Mollusca" rank="species" species="obsoletus">
<emphasis id="B96BD427EE13FFC78E64648E3AB76BA0" box="[303,429,687,708]" italics="true" pageId="37" pageNumber="484">I. obsoletus</emphasis>
</taxonomicName>
the left arm of the visceral loop branches from
<taxonomicName id="4C1F73B6EE13FFC78ED964EC39316B87" baseAuthorityName="Robson" baseAuthorityYear="1920" box="[402,555,717,739]" genus="The" pageId="37" pageNumber="502" rank="species" species="posterior">the posterior</taxonomicName>
osphradial nerve near the osphradium; in other nassariids, the visceral loop arises much closer to the circum-oesophageal nerve ring (Brown) or from the nerve ring itself (Fretter &amp; Graham). Three or five pairs of buccal nerves innervate the proboscis (Brown). The propodial ganglia may be lacking; a single visceral ganglion with two accessory ganglia may be present (Brown).
</paragraph>
</subSubSection>
</treatment>
</document>
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