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<document id="D0365DCBE8130B362B1983522BB953E3" 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="03B6B923EE23FFF48FB3606C3E0B6C2D" 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="Littorina littorea LINNAEUS 1758" docType="treatment" docVersion="4" lastPageNumber="469" masterDocId="FF8FC15BEE36FFE28F4B66213B1A6964" masterDocTitle="Refining molluscan characters: morphology, character coding and a phylogeny of the Caenogastropoda" masterLastPageNumber="554" masterPageNumber="447" pageNumber="468" 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="310C86B4CEED49832CDC9CD0EFF7A156">Refining molluscan characters: morphology, character coding and a phylogeny of the Caenogastropoda</mods:title>
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<treatment id="03B6B923EE23FFF48FB3606C3E0B6C2D" ID-DOI="http://doi.org/10.5281/zenodo.5490932" ID-Zenodo-Dep="5490932" LSID="urn:lsid:plazi:treatment:03B6B923EE23FFF48FB3606C3E0B6C2D" httpUri="http://treatment.plazi.org/id/03B6B923EE23FFF48FB3606C3E0B6C2D" lastPageId="22" lastPageNumber="469" pageId="21" pageNumber="468">
<subSubSection id="C3055BBEEE23FFF78FB3606C39BF6F00" box="[248,677,1612,1636]" pageId="21" pageNumber="468" type="nomenclature">
<paragraph id="8BA00835EE23FFF78FB3606C39BF6F00" blockId="21.[248,677,1580,1636]" box="[248,677,1612,1636]" pageId="21" pageNumber="468">
<heading id="D0E8BF59EE23FFF78FB3606C39BF6F00" box="[248,677,1612,1636]" centered="true" fontSize="9" level="2" pageId="21" pageNumber="468" reason="2">
<taxonomicName id="4C1F73B6EE23FFF78FB3606C39BF6F00" ID-CoL="3VNPR" authority="LINNAEUS 1758" authorityName="LINNAEUS" authorityYear="1758" box="[248,677,1612,1636]" class="Gastropoda" family="Littorinidae" genus="Littorina" kingdom="Animalia" order="Littorinimorpha" pageId="21" pageNumber="468" phylum="Mollusca" rank="species" species="littorea">
<emphasis id="B96BD427EE23FFF78FB3606C3AF26F06" box="[248,488,1613,1636]" italics="true" pageId="21" pageNumber="468">LITTORINA LITTOREA</emphasis>
LINNAEUS 1758
</taxonomicName>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="C3055BBEEE23FFF48FD060553E0B6C2D" lastPageId="22" lastPageNumber="469" pageId="21" pageNumber="468" type="materials_examined">
<paragraph id="8BA00835EE23FFF78FD060553A766FED" blockId="21.[155,678,1652,1705]" box="[155,364,1652,1673]" pageId="21" pageNumber="468">
<emphasis id="B96BD427EE23FFF78FD060553A766FED" box="[155,364,1652,1673]" italics="true" pageId="21" pageNumber="468">Material examined</emphasis>
</paragraph>
<paragraph id="8BA00835EE23FFF78FD060B239BC6FCD" blockId="21.[155,678,1652,1705]" box="[155,678,1683,1705]" pageId="21" pageNumber="468">
Woods Hole,
<collectingRegion id="49DBC6D7EE23FFF78E7860B23AC66FCD" box="[307,476,1683,1705]" country="United States of America" name="Massachusetts" pageId="21" pageNumber="468">Massachusetts</collectingRegion>
(
<collectionCode id="ED0E90F0EE23FFF78EA760B239266FCC" box="[492,572,1683,1704]" 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="21" pageNumber="468" type="Museum">USNM</collectionCode>
890939).
</paragraph>
<paragraph id="8BA00835EE23FFF78FD060DF39296E70" blockId="21.[155,771,1790,1905]" box="[155,563,1790,1812]" pageId="21" pageNumber="468">
<emphasis id="B96BD427EE23FFF78FD060DF39296E70" box="[155,563,1790,1812]" italics="true" pageId="21" pageNumber="468">External anatomy and mantle cavity</emphasis>
</paragraph>
<paragraph id="8BA00835EE23FFF78FD0613C39126E15" blockId="21.[155,771,1790,1905]" pageId="21" pageNumber="468">Pedal gland opening to anterior margin of propodium. Operculum present. Mantle margin smooth. Hypobranchial gland well developed.</paragraph>
<paragraph id="8BA00835EE23FFF78C7866E53F0069BD" blockId="21.[819,1435,196,494]" box="[819,1050,196,217]" pageId="21" pageNumber="468">
<emphasis id="B96BD427EE23FFF78C7866E53F0069BD" box="[819,1050,196,217]" italics="true" pageId="21" pageNumber="468">Reproductive System</emphasis>
</paragraph>
<paragraph id="8BA00835EE23FFF78C7866C23F586817" blockId="21.[819,1435,196,494]" pageId="21" pageNumber="468">Renal oviduct functioning as seminal receptacle, storing orientated sperm. Gonopericardial canal absent. Oviduct opening to glandular gonoduct at base of mantle cavity. Pallial oviduct closed along entire length. Anterior bursa present.</paragraph>
<paragraph id="8BA00835EE23FFF78C00675D3F88688A" blockId="21.[819,1435,196,494]" pageId="21" pageNumber="468">Proximal vas deferens convoluted, forming seminal vesicle. Prostate open to mantle cavity. Open seminal groove extending from prostate to tip of penis lying behind right cephalic tentacle.</paragraph>
<paragraph id="8BA00835EE23FFF78C7864623F1B6B3D" blockId="21.[819,1435,579,1184]" box="[819,1025,579,601]" pageId="21" pageNumber="468">
<emphasis id="B96BD427EE23FFF78C7864623F1B6B3D" box="[819,1025,579,601]" italics="true" pageId="21" pageNumber="468">Alimentary system</emphasis>
</paragraph>
<paragraph id="8BA00835EE23FFF78C7864423F5B6DC4" blockId="21.[819,1435,579,1184]" pageId="21" pageNumber="468">
<emphasis id="B96BD427EE23FFF78C786442388B6B1C" box="[819,913,611,632]" italics="true" pageId="21" pageNumber="468">Foregut.</emphasis>
Radula taenioglossate. Paired cartilages supporting odontophore. Subradular membrane almost completely covering odontophore. Weakly glandular, narrow papilla, extending forward ventral to radular ribbon (
<figureCitation id="132414B0EE23FFF78CD964FC38F66B96" box="[914,1004,733,755]" captionStart="Figure 4" captionStartId="46.[142,220,1755,1774]" captionTargetBox="[274,1290,197,1722]" captionTargetId="figure-144@46.[274,1291,197,1722]" captionTargetPageId="46" captionText="Figure 4. Morphology of odontophore; subradular organ (indicated by arrows). Except when noted, view is from the side and ventral is to the right. (A) Theodoxus fluviatilis, ventral view. Subradular organ absent. Scale bar = 100 Mm. (B) Neocyclotus dysoni ambiguum. Scale bar = 1 mm. (C) Marisa cornuarietis. Scale bar = 1 mm. (D) Lampanella minima. Scale bar = 100 Mm. (E) Littorina littorea. Scale bar = 100 Mm. (F) Neverita duplicata. Scale bar = 100 Mm. (G) Cypraea robertsi. Subradular organ absent. Scale bar = 1 mm. (H) Panarona clavatula. Ventral is down. Scale bar = 100 Mm." figureDoi="http://doi.org/10.5281/zenodo.5437615" httpUri="https://zenodo.org/record/5437615/files/figure.png" pageId="21" pageNumber="468">Figs 4E</figureCitation>
,
<figureCitation id="132414B0EE23FFF78CB564FD3F046B96" box="[1022,1054,732,754]" captionStart="Figure 5" captionStartId="48.[142,220,1696,1715]" captionTargetBox="[293,1272,197,1664]" captionTargetId="figure-150@48.[293,1272,197,1664]" captionTargetPageId="48" captionText="Figure 5. Histology of subradular organ. (A) Neocyclotus dysoni ambiguum. Transverse section. Note bi-layered jaw. Scale bar = 0.5 mm. (B) Marisa cornuarietis. Sagittal section. Note bi-layered jaw. Scale bar = 0.5 mm. (C) Lampanella minima. Transverse section. Note jaw with exposed rods. Scale bar = 1 mm. (D) Strombus mutabilis. Transverse section. Scale bar = 0.5 mm. (E) Crepidula plana. Transverse section. Scale bar = 1 mm. (F) Littorina littorea. Sagittal section. Scale bar = 1 mm. (G) Neverita duplicata. Transverse section. Scale bar = 0.5 mm. (H) Panarona clavatula. Transverse section. Scale bar = 1 mm. Abbreviations: c., radular cartilage; j, jaw; mo, mouth; r, radula; srm, subradular membrane; sro, subradular organ." figureDoi="http://doi.org/10.5281/zenodo.5437617" httpUri="https://zenodo.org/record/5437617/files/figure.png" pageId="21" pageNumber="468">5F</figureCitation>
, sro). Dorsal surface of papilla lined with thin layer of cuticle. Buccal cavity lined with cuticle. Glandular mid-ventral fold (
<figureCitation id="132414B0EE23FFF78A50653B3E776A4B" box="[1307,1389,794,816]" captionStart="Figure 8" captionStartId="53.[155,233,1725,1744]" captionTargetBox="[296,1293,197,1693]" captionTargetId="figure-144@53.[296,1294,197,1693]" captionTargetPageId="53" captionText="Figure 8. Histology of anterior oesophagus; ventral fold. (A) Macleaniella moskalevi. Scale bar = 0.5 mm. (B) Theodoxus fluviatilis. Scale bar = 0.5 mm. (C) Neocyclotus dysoni ambiguum. Scale bar = 0.5 mm. (D) Lampanella minima. Scale bar = 1 mm. (E) Strombus mutabilis. Scale bar = 0.5 mm. (F) Littorina littorea. Scale bar = 0.5 mm. (G) Cypraea robertsi. Note that ventral fold is offset to the right. Scale bar = 0.5 mm. Abbreviations: bp, buccal pouch; bg, buccal ganglion; c., radular cartilage; ceg, cerebral ganglion; df, dorsal fold; e.g. oesophageal gland; r, radula; sgd, salivary gland duct; sgl, salivary gland; vf, ventral fold; vlf, ventro-lateral fold." figureDoi="http://doi.org/10.5281/zenodo.5437623" httpUri="https://zenodo.org/record/5437623/files/figure.png" pageId="21" pageNumber="468">Fig. 8F</figureCitation>
, vf) beginning in buccal cavity, extending posteriorly to buccal pouches, then diminishing. Mid-ventral fold not continuing into mid-oesophagus. Buccal pouches opening to posterior buccal cavity and anterior oesophagus. Anterior ducts of buccal pouches lined with highly glandular, acidophilic epithelium, expanding dorsally into buccal pouches. Remainder of pouch epithelium strongly basophilic. Apertures of pouches bounded by prominent ventro-lateral folds (
<figureCitation id="132414B0EE23FFF78BE8620F3FE06D27" box="[1187,1274,1070,1092]" captionStart="Figure 9" captionStartId="54.[142,220,1725,1744]" captionTargetBox="[283,1280,197,1693]" captionTargetId="figure-140@54.[283,1281,197,1693]" captionTargetPageId="54" captionText="Figure 9. Histology of anterior oesophagus; ventro-lateral folds. (A) Macleaniella moskalevi. Scale bar = 0.5 mm. (B) Theodoxus fluviatilis. Scale bar = 0.5 mm. (C) Neocyclotus dysoni ambiguum. Scale bar = 0.5 mm. (D) Lampanella minima. Scale bar = 1 mm. (E) Strombus mutabilis. Scale bar = 0.5 mm. (F) Crepidula plana. Scale bar = 1 mm. (G) Bithynia tentaculata. Scale bar = 1 mm. (H) Littorina littorea. Scale bar = 0.5 mm. Abbreviations: bp, buccal pouch; bg, buccal ganglion; ca, radular cartilage; ceg, cerebral ganglion; df, dorsal fold; e, oesophagus; r, radula; sgl, salivary gland; vf, ventral fold; vlf, ventro-lateral fold." figureDoi="http://doi.org/10.5281/zenodo.5437625" httpUri="https://zenodo.org/record/5437625/files/figure.png" pageId="21" pageNumber="468">Fig. 9H</figureCitation>
, vlf). Oesophageal gland present within mid-oesophagus, forming transverse septae of glandular tissue partially occluding oesophageal lumen.
</paragraph>
<paragraph id="8BA00835EE23FFF78C78629938DC6E15" blockId="21.[819,1435,1208,1905]" pageId="21" pageNumber="468">
<emphasis id="B96BD427EE23FFF78C78629938946DA9" box="[819,910,1208,1229]" italics="true" pageId="21" pageNumber="468">Midgut.</emphasis>
Oesophagus (
<figureCitation id="132414B0EE23FFF78B7F62993FB56DA9" box="[1076,1199,1208,1230]" captionStart="Figure 15" captionStartId="64.[142,220,1725,1744]" captionTargetBox="[283,1268,197,1693]" captionTargetId="figure-125@64.[283,1281,197,1693]" captionTargetPageId="64" captionText="Figure 15. Midgut morphology. White arrows indicate direction of ciliary currents. (A) Bithynia tentaculata, dorsal view. Distal end of gastric chamber obscured in scanning electron micrograph, indicated by dotted line in diagram on right. (B) Littorina littorea, dorsal view. Stippling indicates ciliary tract within gastric chamber extending from oesophageal aperture. Scale bar = 1 mm. Abbreviations: c, caecum; ce, caecal extension; cf, ciliated fold; cs, ciliated strip; ctr, ciliary tract; dgd, digestive gland duct; e, oesophagus; gp, glandular pad; gs, gastric shield; sa, sorting area; ss, style sac; t1, major typhlosole; t2, minor typhlosole." figureDoi="http://doi.org/10.5281/zenodo.5437639" httpUri="https://zenodo.org/record/5437639/files/figure.png" pageId="21" pageNumber="468">Fig. 15B, e</figureCitation>
) opening near proximal end of minor typhlosole (t2); mucus-secreting pouch at base of minor typhlosole absent. Three ducts of digestive gland (dgd) and surrounding ciliary sorting area (sa) lying at proximal end of major typhlosole (t2). Ciliary currents in sorting area flowing anteriorly toward intestinal groove. Gastric shield (gs) ventrolateral on right. Gastric chamber cuticularized. Ciliated fold (cf) bearing short, dense cilia emerging near minor typhlosole, running posteriorly to left of oesophageal aperture. Ciliated fold bordering grooved tract bearing long cilia. Grooved tract emerging from under lip of ciliated ridge near oesophageal aperture, paralleling ridge posteriorly. Grooved tract curving around posterior periphery of gastric chamber, extending short distance anteriorly along right wall. Currents within ciliary tract flowing posteriorly. Low, ciliated ridge (cs) extending posteriorly along left side of gastric shield. Currents over ciliated ridge flowing posteriorly and towards right side. Crystalline style absent. Raised tract of cilia on major typhlosole absent. Currents flowing clockwise within style sac when viewed from behind.
</paragraph>
<paragraph id="8BA00835EE20FFF48FC566E53A1E6873" blockId="22.[142,757,196,279]" pageId="22" pageNumber="469">
<emphasis id="B96BD427EE20FFF48FC566E53BEF69BD" box="[142,245,196,217]" italics="true" pageId="22" pageNumber="469">Hindgut.</emphasis>
Intestine emerging from style sac, curving posteriorly around rear wall of kidney before turning anteriorly.
</paragraph>
<paragraph id="8BA00835EE20FFF48FC5677C3A806817" blockId="22.[142,758,349,494]" box="[142,410,349,371]" pageId="22" pageNumber="469">
<emphasis id="B96BD427EE20FFF48FC5677C3A806817" box="[142,410,349,371]" italics="true" pageId="22" pageNumber="469">Reno-pericardial system</emphasis>
</paragraph>
<paragraph id="8BA00835EE20FFF48FC5675D39C5688A" blockId="22.[142,758,349,494]" pageId="22" pageNumber="469">Afferent renal vessel entering kidney floor anteriorly, extending dorsally and ramifying to supply excretory tissue. Nephridial gland present. Branch from afferent renal vessel supplying nephridial gland absent.</paragraph>
<paragraph id="8BA00835EE20FFF48FC5641539296B2D" blockId="22.[142,758,564,831]" box="[142,563,564,585]" pageId="22" pageNumber="469">
<emphasis id="B96BD427EE20FFF48FC5641539296B2D" box="[142,563,564,585]" italics="true" pageId="22" pageNumber="469">Nervous system and sensory structure</emphasis>
</paragraph>
<paragraph id="8BA00835EE20FFF48FC56472395C6A5B" blockId="22.[142,758,564,831]" pageId="22" pageNumber="469">Nervous system epiathroid, left dialyneurous. Nerve ring lying just behind base of buccal mass. Tentacular nerve bifid. Metapodial commissure present. Visceral ganglia paired. Statocysts with single statoliths present postero-dorsally on pedal ganglia, slightly asymmetrical. Small eyes present at lateral bases of cephalic tentacles. Osphradium comprising long, thin, ciliated ridge along base of ctenidium.</paragraph>
<paragraph id="8BA00835EE20FFF48FC565A43BEA6AFE" blockId="22.[142,758,901,1659]" box="[142,240,901,922]" pageId="22" pageNumber="469">
<emphasis id="B96BD427EE20FFF48FC565A43BEA6AFE" box="[142,240,901,922]" italics="true" pageId="22" pageNumber="469">Remarks</emphasis>
</paragraph>
<paragraph id="8BA00835EE20FFF48FC565853A516DAA" blockId="22.[142,758,901,1659]" pageId="22" pageNumber="469">
The anatomy of this species has been well established in several published descriptions, most notably a thorough account by
<bibRefCitation id="EF8E75C4EE20FFF48E1C65C3399E6A93" author="Fretter V &amp; Graham A" box="[343,644,993,1015]" pageId="22" pageNumber="469" 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>
. Previous studies include those of
<bibRefCitation id="EF8E75C4EE20FFF48EE36221392F6D72" author="Ankel WE" box="[424,565,1024,1046]" pageId="22" pageNumber="469" pagination="1 - 240" refId="ref71937" refString="Ankel WE. 1936. Prosobranchia. In: Grimpe G, Wagler E, eds. Die Tierwelt der Nord- und Ostsee. Leipzig: Akademische Verlagsgesellschaft, 1 - 240." type="book chapter" year="1936">Ankel (1936</bibRefCitation>
,
<bibRefCitation id="EF8E75C4EE20FFF48D08622139666D72" author="Ankel WE" box="[579,636,1024,1046]" pageId="22" pageNumber="469" pagination="317 - 333" refId="ref71970" refString="Ankel WE. 1937. Wie frisst Littorina? Senckenbergiana 19: 317 - 333." type="journal article" year="1937">1937</bibRefCitation>
), who provided information on foregut anatomy and the nervous sytsem, and
<bibRefCitation id="EF8E75C4EE20FFF48E79621C39196D37" author="Johansson J" box="[306,515,1085,1107]" pageId="22" pageNumber="469" pagination="288 - 396" refId="ref76119" refString="Johansson J. 1939. Anatomische Studien uber die Gastropodenfamilien Rissoidae und Littorinidae. Zoologiska Bidrag Fran Uppsala 18: 288 - 396." type="journal article" year="1939">Johansson (1939)</bibRefCitation>
who also provided details on foregut anatomy. Lacking from these accounts are descriptions of the unique, papilla-like subradular organ, and the ventral folding of the anterior oesophagus.
</paragraph>
<paragraph id="8BA00835EE20FFF48FEC62F639776F7B" blockId="22.[142,758,901,1659]" pageId="22" pageNumber="469">
<bibRefCitation id="EF8E75C4EE20FFF48FEC62F63A6D6D89" author="Johansson J" box="[167,375,1239,1261]" pageId="22" pageNumber="469" pagination="288 - 396" refId="ref76119" refString="Johansson J. 1939. Anatomische Studien uber die Gastropodenfamilien Rissoidae und Littorinidae. Zoologiska Bidrag Fran Uppsala 18: 288 - 396." type="journal article" year="1939">Johansson (1939)</bibRefCitation>
and
<bibRefCitation id="EF8E75C4EE20FFF48E8962F639606D89" author="Graham A" box="[450,634,1239,1261]" pageId="22" pageNumber="469" 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>
described midgut anatomy for this species. The present description agrees with these on most major points. However, there are several discrepancies. Both Johansson and Graham described only two ciliary tracts leading posteriorly into the gastric chamber; one bounding the oesophagus to the left and a second passing to the left of the gastric shield. In addition, Graham found a mucus-secreting pouch at the base of the minor typhlosole; ciliary currents were described as flowing anteriorly to the left of the gastric shield.
</paragraph>
<paragraph id="8BA00835EE20FFF48FEC600939FA6F1F" blockId="22.[142,758,901,1659]" pageId="22" pageNumber="469">
The metapodial commissure has been reported as present (
<bibRefCitation id="EF8E75C4EE20FFF48FBF606639096F39" author="Fretter V &amp; Graham A" box="[244,531,1607,1629]" pageId="22" pageNumber="469" 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="EF8E75C4EE20FFF48D5460663BDF6F1F" author="Marcus E &amp; Marcus E" pageId="22" pageNumber="469" pagination="5 - 105" refId="ref77110" refString="Marcus E, Marcus E. 1963. Mesogastropoden von der Kuste Sao Paulos. Akademie der Wissenschaften und der Literatur, Mathematisch-Naturwissenschaftliche Klasse 1963 (1): 5 - 105." type="journal article" year="1963">Marcus &amp; Marcus, 1963</bibRefCitation>
) or absent (
<bibRefCitation id="EF8E75C4EE20FFF48E1A60443AEF6F1F" author="Simroth H" box="[337,501,1637,1659]" pageId="22" pageNumber="469" refId="ref79936" refString="Simroth H. 1896 - 1907. Gastropoda prosobranchia. H. G. Bronn's Klassen und Ordnungen Des Thierreichs, Wissenschaftlich Dargestellt in Wort und Bild, Vol. 3. Mollusca, Abteilung II. Leipzig: CF Winter'sche Verlagshandlung." type="book" year="1896">Simroth, 1896</bibRefCitation>
1907;
<bibRefCitation id="EF8E75C4EE20FFF48D0C604439CE6F1F" author="Ankel WE" box="[583,724,1637,1659]" pageId="22" pageNumber="469" pagination="1 - 240" refId="ref71937" refString="Ankel WE. 1936. Prosobranchia. In: Grimpe G, Wagler E, eds. Die Tierwelt der Nord- und Ostsee. Leipzig: Akademische Verlagsgesellschaft, 1 - 240." type="book chapter" year="1936">Ankel, 1936</bibRefCitation>
).
</paragraph>
<paragraph id="8BA00835EE20FFF48FC560E33A1F6FB3" blockId="22.[142,758,1730,1905]" box="[142,261,1730,1751]" pageId="22" pageNumber="469">
<emphasis id="B96BD427EE20FFF48FC560E33A1F6FB3" box="[142,261,1730,1751]" italics="true" pageId="22" pageNumber="469">Discussion</emphasis>
</paragraph>
<paragraph id="8BA00835EE20FFF48FC560C13F716831" blockId="22.[142,758,1730,1905]" lastBlockId="22.[806,1422,196,1353]" pageId="22" pageNumber="469">
The basic layout of the reproductive system and the foregut of littorinids is largely congruent with
<taxonomicName id="4C1F73B6EE20FFF48D8760DE3A376E56" authorityName="LINNAEUS" authorityYear="1758" class="Gastropoda" family="Littorinidae" genus="Littorina" kingdom="Animalia" order="Littorinimorpha" pageId="22" pageNumber="469" phylum="Mollusca" rank="species" species="littorea">
<emphasis id="B96BD427EE20FFF48D8760DE3A376E56" italics="true" pageId="22" pageNumber="469">Littorina littorea</emphasis>
</taxonomicName>
. Variable features of reproductive anatomy include the presence or absence of a gonopericardial canal, the looping patterns of the gladular oviduct, an open or closed prostate and seminal groove, and the morphology of the penis and accessory glands (e.g.
<bibRefCitation id="EF8E75C4EE20FFF48B6767203FA26872" author="Linke O" box="[1068,1208,257,279]" pageId="22" pageNumber="469" pagination="2 - 60" refId="ref76836" refString="Linke O. 1933. Morphologie und Physiologie des Genitalapparates der Nordseelittorinen. Wissenschaftliche Meeresuntersuchungen 19: 2 - 60." type="journal article" year="1933">Linke, 1933</bibRefCitation>
;
<bibRefCitation id="EF8E75C4EE20FFF48B8C67203E906872" author="Johansson J" box="[1223,1418,257,279]" pageId="22" pageNumber="469" pagination="288 - 396" refId="ref76119" refString="Johansson J. 1939. Anatomische Studien uber die Gastropodenfamilien Rissoidae und Littorinidae. Zoologiska Bidrag Fran Uppsala 18: 288 - 396." type="journal article" year="1939">Johansson, 1939</bibRefCitation>
;
<bibRefCitation id="EF8E75C4EE20FFF48C6D67013F486852" author="Fretter V &amp; Graham A" box="[806,1106,288,310]" pageId="22" pageNumber="469" 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="EF8E75C4EE20FFF48B2867013E906852" author="Marcus E &amp; Marcus E" box="[1123,1418,288,310]" pageId="22" pageNumber="469" pagination="5 - 105" refId="ref77110" refString="Marcus E, Marcus E. 1963. Mesogastropoden von der Kuste Sao Paulos. Akademie der Wissenschaften und der Literatur, Mathematisch-Naturwissenschaftliche Klasse 1963 (1): 5 - 105." type="journal article" year="1963">Marcus &amp; Marcus, 1963</bibRefCitation>
;
<bibRefCitation id="EF8E75C4EE20FFF48C6D671E38CC6830" author="Bingham FO" box="[806,982,319,341]" pageId="22" pageNumber="469" pagination="8 - 10" refId="ref72489" refString="Bingham FO. 1972. Several aspects of the reproductive biology of Littorina irrorata. The Nautilus 86: 8 - 10." type="journal article" year="1972">Bingham, 1972</bibRefCitation>
;
<bibRefCitation id="EF8E75C4EE20FFF48CA9671E3F7A6831" author="Reid DG" box="[994,1120,319,341]" pageId="22" pageNumber="469" refId="ref79067" refString="Reid DG. 1986. The littorinid molluscs of mangrove forests in the Indo-Pacific region. London: British Museum (Natural History)." type="book" year="1986">Reid, 1986</bibRefCitation>
).
</paragraph>
<paragraph id="8BA00835EE20FFF48C74677C38B66D72" blockId="22.[806,1422,196,1353]" pageId="22" pageNumber="469">
All littorinids described possess an elongate, cuticularized gastric chamber, with at least one ciliary tract (e.g.
<bibRefCitation id="EF8E75C4EE20FFF48CED67BA3FD468D5" author="Marcus E &amp; Marcus E" box="[934,1230,411,433]" pageId="22" pageNumber="469" pagination="5 - 105" refId="ref77110" refString="Marcus E, Marcus E. 1963. Mesogastropoden von der Kuste Sao Paulos. Akademie der Wissenschaften und der Literatur, Mathematisch-Naturwissenschaftliche Klasse 1963 (1): 5 - 105." type="journal article" year="1963">Marcus &amp; Marcus, 1963</bibRefCitation>
;
<bibRefCitation id="EF8E75C4EE20FFF48BAB67BA3E9868D4" author="Morton JE" box="[1248,1410,410,432]" pageId="22" pageNumber="469" pagination="1 - 15" refId="ref77805" refString="Morton JE. 1975. Form and habit in some small gastropods of New Zealand boulder beaches. The Veliger 18: 1 - 15." type="journal article" year="1975">Morton, 1975</bibRefCitation>
); length of the gastric chamber is variable and is not correlated with habitat (
<bibRefCitation id="EF8E75C4EE20FFF48B0367F93FD3688A" author="Reid DG" box="[1096,1225,472,494]" pageId="22" pageNumber="469" refId="ref79067" refString="Reid DG. 1986. The littorinid molluscs of mangrove forests in the Indo-Pacific region. London: British Museum (Natural History)." type="book" year="1986">Reid, 1986</bibRefCitation>
). The number of ciliary tracts and digestive gland ducts and the size of the sorting area are also variable (e.g.
<bibRefCitation id="EF8E75C4EE20FFF48A5E643738D36B2E" author="Marcus E &amp; Marcus E" pageId="22" pageNumber="469" pagination="5 - 105" refId="ref77110" refString="Marcus E, Marcus E. 1963. Mesogastropoden von der Kuste Sao Paulos. Akademie der Wissenschaften und der Literatur, Mathematisch-Naturwissenschaftliche Klasse 1963 (1): 5 - 105." type="journal article" year="1963">Marcus &amp; Marcus, 1963</bibRefCitation>
;
<bibRefCitation id="EF8E75C4EE20FFF48C9664153F7B6B2E" author="Reid DG" box="[989,1121,564,586]" pageId="22" pageNumber="469" refId="ref79067" refString="Reid DG. 1986. The littorinid molluscs of mangrove forests in the Indo-Pacific region. London: British Museum (Natural History)." type="book" year="1986">Reid, 1986</bibRefCitation>
). Despite inconsistencies between the present and previous descriptions as summarized above, the present results are congruent with investigations of
<taxonomicName id="4C1F73B6EE20FFF48B6D64B03E116BC2" baseAuthorityName="Linnaeus" baseAuthorityYear="1758" box="[1062,1291,657,678]" class="Gastropoda" family="Littorinidae" genus="Tectarius" kingdom="Animalia" order="Littorinimorpha" pageId="22" pageNumber="469" phylum="Mollusca" rank="species" species="muricatus">
<emphasis id="B96BD427EE20FFF48B6D64B03E116BC2" box="[1062,1291,657,678]" italics="true" pageId="22" pageNumber="469">Tectarius muricatus</emphasis>
</taxonomicName>
(pers. obs.) and
<taxonomicName id="4C1F73B6EE20FFF48C2E648E3E426BA0" box="[869,1368,687,708]" class="Gastropoda" family="Littorinidae" genus="Littorina" kingdom="Animalia" order="Littorinimorpha" pageId="22" pageNumber="469" phylum="Mollusca" rank="subSpecies" species="unifasciata" subGenus="Austrolittorina" subSpecies="antipoda">
<emphasis id="B96BD427EE20FFF48C2E648E38666BA0" box="[869,892,687,708]" italics="true" pageId="22" pageNumber="469">L.</emphasis>
(
<emphasis id="B96BD427EE20FFF48CD3648E3F586BA0" box="[920,1090,687,708]" italics="true" pageId="22" pageNumber="469">Austrolittorina</emphasis>
)
<emphasis id="B96BD427EE20FFF48B15648E3E426BA0" box="[1118,1368,687,708]" italics="true" pageId="22" pageNumber="469">unifasciata antipoda</emphasis>
</taxonomicName>
(as
<taxonomicName id="4C1F73B6EE20FFF48C6D64EC3F1A6B86" authorityName="Finlay" authorityYear="1930" box="[806,1024,717,738]" class="Gastropoda" family="Littorinidae" genus="Melarhaphe" kingdom="Animalia" order="Littorinimorpha" pageId="22" pageNumber="469" phylum="Mollusca" rank="species" species="oliveri">
<emphasis id="B96BD427EE20FFF48C6D64EC3F1A6B86" box="[806,1024,717,738]" italics="true" pageId="22" pageNumber="469">Melarhaphe oliveri</emphasis>
</taxonomicName>
) (
<bibRefCitation id="EF8E75C4EE20FFF48B5164EC3FA26B87" author="Morton JE" box="[1050,1208,717,739]" pageId="22" pageNumber="469" pagination="1 - 15" refId="ref77805" refString="Morton JE. 1975. Form and habit in some small gastropods of New Zealand boulder beaches. The Veliger 18: 1 - 15." type="journal article" year="1975">Morton, 1975</bibRefCitation>
). All three species possess three ciliary tracts; one extending from the oesophageal aperture and curving around the base of the gastric chamber with two flanking tracts. Similar to
<bibRefCitation id="EF8E75C4EE20FFF48CED65693F436A3A" author="Graham A" box="[934,1113,840,862]" pageId="22" pageNumber="469" 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>
,
<bibRefCitation id="EF8E75C4EE20FFF48B2365693E946A3A" author="Marcus E &amp; Marcus E" box="[1128,1422,840,862]" pageId="22" pageNumber="469" pagination="5 - 105" refId="ref77110" refString="Marcus E, Marcus E. 1963. Mesogastropoden von der Kuste Sao Paulos. Akademie der Wissenschaften und der Literatur, Mathematisch-Naturwissenschaftliche Klasse 1963 (1): 5 - 105." type="journal article" year="1963">Marcus &amp; Marcus (1963)</bibRefCitation>
documented a mucus-secreting pouch at the proximal end of the style sac in
<taxonomicName id="4C1F73B6EE20FFF48B1765A73FEA6AFE" baseAuthorityName="Lamarck" baseAuthorityYear="1822" box="[1116,1264,901,923]" class="Gastropoda" family="Littorinidae" genus="Littorina" kingdom="Animalia" order="Littorinimorpha" pageId="22" pageNumber="469" phylum="Mollusca" rank="species" species="angulifera">
<emphasis id="B96BD427EE20FFF48B1765A73FEA6AFE" box="[1116,1264,901,923]" italics="true" pageId="22" pageNumber="469">L. angulifera</emphasis>
</taxonomicName>
.
<taxonomicName id="4C1F73B6EE20FFF48BB565A738456ADE" authorityName="LINNAEUS" authorityYear="1758" class="Gastropoda" family="Littorinidae" genus="Littorina" kingdom="Animalia" order="Littorinimorpha" pageId="22" pageNumber="469" phylum="Mollusca" rank="species" species="littorea">
<emphasis id="B96BD427EE20FFF48BB565A738456ADE" italics="true" pageId="22" pageNumber="469">Littorina littorea</emphasis>
</taxonomicName>
is intriguing because it lacks a ciliary tract on the major typhlosole; the tract is present in another non-style-bearing littorinid,
<taxonomicName id="4C1F73B6EE20FFF48BB565C33E946A93" baseAuthorityName="Linnaeus" baseAuthorityYear="1758" box="[1278,1422,994,1015]" class="Gastropoda" family="Littorinidae" genus="Tectarius" kingdom="Animalia" order="Littorinimorpha" pageId="22" pageNumber="469" phylum="Mollusca" rank="species" species="muricatus">
<emphasis id="B96BD427EE20FFF48BB565C33E946A93" box="[1278,1422,994,1015]" italics="true" pageId="22" pageNumber="469">T. muricatus</emphasis>
</taxonomicName>
(pers. obs.).
</paragraph>
<paragraph id="8BA00835EE20FFF48C74623E3E0B6C2D" blockId="22.[806,1422,196,1353]" pageId="22" pageNumber="469">
Although much of the detailed information regarding the littorinid nervous system concerns
<taxonomicName id="4C1F73B6EE20FFF48A6C621F38626D15" authorityName="LINNAEUS" authorityYear="1758" class="Gastropoda" family="Littorinidae" genus="Littorina" kingdom="Animalia" order="Littorinimorpha" pageId="22" pageNumber="469" phylum="Mollusca" rank="species" species="littorea">
<emphasis id="B96BD427EE20FFF48A6C621F38626D15" italics="true" pageId="22" pageNumber="469">Littorina littorea</emphasis>
</taxonomicName>
(
<bibRefCitation id="EF8E75C4EE20FFF48CC6627D3F286D15" author="Bouvier E-L" box="[909,1074,1116,1138]" pageId="22" pageNumber="469" refId="ref72611" refString="Bouvier E-L. 1887. Systeme nerveux, morphologie generale et classification des gasteropodes prosobranches. Academie de Paris." type="book" year="1887">Bouvier, 1887</bibRefCitation>
;
<bibRefCitation id="EF8E75C4EE20FFF48B0F627D3E126D15" author="Johansson J" box="[1092,1288,1116,1138]" pageId="22" pageNumber="469" pagination="288 - 396" refId="ref76119" refString="Johansson J. 1939. Anatomische Studien uber die Gastropodenfamilien Rissoidae und Littorinidae. Zoologiska Bidrag Fran Uppsala 18: 288 - 396." type="journal article" year="1939">Johansson, 1939</bibRefCitation>
;
<bibRefCitation id="EF8E75C4EE20FFF48A51627D38D66DF5" author="Fretter V &amp; Graham A" pageId="22" pageNumber="469" 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>
), additional studies have found no significant differences in organization in other species (
<bibRefCitation id="EF8E75C4EE20FFF48C65629938ED6DA9" author="Johansson J" box="[814,1015,1208,1230]" pageId="22" pageNumber="469" pagination="288 - 396" refId="ref76119" refString="Johansson J. 1939. Anatomische Studien uber die Gastropodenfamilien Rissoidae und Littorinidae. Zoologiska Bidrag Fran Uppsala 18: 288 - 396." type="journal article" year="1939">Johansson, 1939</bibRefCitation>
;
<bibRefCitation id="EF8E75C4EE20FFF48B4662993E586DAA" author="Marcus E &amp; Marcus E" box="[1037,1346,1208,1230]" pageId="22" pageNumber="469" pagination="5 - 105" refId="ref77110" refString="Marcus E, Marcus E. 1963. Mesogastropoden von der Kuste Sao Paulos. Akademie der Wissenschaften und der Literatur, Mathematisch-Naturwissenschaftliche Klasse 1963 (1): 5 - 105." type="journal article" year="1963">Marcus &amp; Marcus, 1963</bibRefCitation>
). One minor exception is the metapodial commissure, which has been reported as present (
<bibRefCitation id="EF8E75C4EE20FFF48BCC62D43E526C6E" author="Johansson J" box="[1159,1352,1269,1291]" pageId="22" pageNumber="469" pagination="288 - 396" refId="ref76119" refString="Johansson J. 1939. Anatomische Studien uber die Gastropodenfamilien Rissoidae und Littorinidae. Zoologiska Bidrag Fran Uppsala 18: 288 - 396." type="journal article" year="1939">Johansson, 1939</bibRefCitation>
;
<bibRefCitation id="EF8E75C4EE20FFF48A1F62D73F3F6C4E" author="Fretter V &amp; Graham A" pageId="22" pageNumber="469" 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="EF8E75C4EE20FFF48B7363353E7A6C4E" author="Marcus E &amp; Marcus E" box="[1080,1376,1300,1322]" pageId="22" pageNumber="469" pagination="5 - 105" refId="ref77110" refString="Marcus E, Marcus E. 1963. Mesogastropoden von der Kuste Sao Paulos. Akademie der Wissenschaften und der Literatur, Mathematisch-Naturwissenschaftliche Klasse 1963 (1): 5 - 105." type="journal article" year="1963">Marcus &amp; Marcus, 1963</bibRefCitation>
) or absent (
<bibRefCitation id="EF8E75C4EE20FFF48CC963123F3C6C2D" author="Simroth H" box="[898,1062,1331,1353]" pageId="22" pageNumber="469" refId="ref79936" refString="Simroth H. 1896 - 1907. Gastropoda prosobranchia. H. G. Bronn's Klassen und Ordnungen Des Thierreichs, Wissenschaftlich Dargestellt in Wort und Bild, Vol. 3. Mollusca, Abteilung II. Leipzig: CF Winter'sche Verlagshandlung." type="book" year="1896">Simroth, 1896</bibRefCitation>
1907;
<bibRefCitation id="EF8E75C4EE20FFF48B3363123E1F6C2D" author="Ankel WE" box="[1144,1285,1331,1353]" pageId="22" pageNumber="469" pagination="1 - 240" refId="ref71937" refString="Ankel WE. 1936. Prosobranchia. In: Grimpe G, Wagler E, eds. Die Tierwelt der Nord- und Ostsee. Leipzig: Akademische Verlagsgesellschaft, 1 - 240." type="book chapter" year="1936">Ankel, 1936</bibRefCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>