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<mods:title id="C0E78C11222A13CD890D3FAFF7FBA59F">A stomach plate to divide them all: a phylogenetic reassessment of the family Tritoniidae (Nudibranchia: Cladobranchia)</mods:title>
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<treatment id="03A6878CFF876500FD42FB5DEBC8FB06" ID-DOI="http://doi.org/10.5281/zenodo.8432427" ID-GBIF-Taxon="213768877" ID-Zenodo-Dep="8432427" LSID="urn:lsid:plazi:treatment:03A6878CFF876500FD42FB5DEBC8FB06" httpUri="http://treatment.plazi.org/id/03A6878CFF876500FD42FB5DEBC8FB06" lastPageNumber="468" pageId="23" pageNumber="468">
<subSubSection id="C3156511FF876500FD42FB5DEC43FA93" pageId="23" pageNumber="468" type="nomenclature">
<paragraph id="8BB0369AFF876500FD42FB5DEC43FA93" blockId="23.[272,604,1241,1304]" pageId="23" pageNumber="468">
<emphasis id="B97BEA88FF876500FD42FB5DEC43FA93" bold="true" pageId="23" pageNumber="468">
Genus
<taxonomicName id="4C0F4D19FF876500FD08FB5DEC3BFB78" ID-CoL="7P8GQ" authority="Vayssiere 1877" authorityName="Vayssiere" authorityYear="1877" box="[346,604,1241,1268]" class="Gastropoda" family="Tritoniidae" genus="Marionia" kingdom="Animalia" order="Nudibranchia" pageId="23" pageNumber="468" phylum="Mollusca" rank="genus">
<emphasis id="B97BEA88FF876500FD08FB5DEFDDFB78" bold="true" box="[346,442,1241,1267]" italics="true" pageId="23" pageNumber="468">Marionia</emphasis>
Vayssière 1877
</taxonomicName>
(
<figureCitation id="13342A1FFF876500FD00FB7AEFFFFA93" box="[338,408,1278,1304]" captionStart="Figure 3" captionStartId="12.[129,194,1533,1557]" captionTargetBox="[321,1281,144,1505]" captionTargetId="figure-116@12.[321,1281,144,1505]" captionTargetPageId="12" captionText="Figure 3. Representative species of genera of Tritoniidae (with exception of Paratritonia). A, Tritonia hombergii (MNCN15.05/93379) (Portaferry, Ireland); B, Candiella plebeia (MNCN15.05/93393) (Portaferry, Ireland); C, Tritoniopsis elegans (MB28-0045061) (Zavora, Mozambique); D, Tritonidoxa capensis (MNCN15.05/93399) (Scoưburg, South Africa); E, Tritoniella belli (Antarctica); F, Marianina rosea (French Polynesia); G, Myrella challengeriana comb.nov.(Bahía Camarones, Argentina); H, Tochuina gigantea (Agnew Pass, USA); I, Tritonicula khaleesi (MZUSP113165) (Fortaleza, Brazil); J, Marionia cucullata (Golfo Nuevo, Argentina); (K) Marioniopsis levis 1 (MNHNC/ MB28-004895) (Nacala, Mozambique). Photo A and B, courtesy of B. Picton; photo C and K courtesy of Y.Tibiriça; photo D courtesy of V. Fraser; photo E courtesy of M. Caballer; photo F courtesy of G. Rouse; photo G and J, courtesy of N. Baưini; photo H courtesy of S. Thiebaud; photo I by F. Silva." figureDoi="http://doi.org/10.5281/zenodo.8432399" httpUri="https://zenodo.org/record/8432399/files/figure.png" pageId="23" pageNumber="468">Figs 3J</figureCitation>
,
<figureCitation id="13342A1FFF876500FDF0FB7AEFA4FA93" box="[418,451,1278,1304]" captionStart="Figure 4" captionStartId="13.[113,178,1528,1552]" captionTargetBox="[309,1263,147,1498]" captionTargetId="figure-134@13.[306,1266,144,1501]" captionTargetPageId="13" captionText="Figure 4. Scanning electron micrographs of radulae of analysed key species of Tritoniinae and Marioniinae subfam. nov.: A, tricuspid rachidian teeth and differentiated first lateral of Tritonia hombergii (ZMBN 83471); B, tricuspid rachidian teeth and differentiated first lateral with single accessory denticle (circled) of Candiella plebeia (MNCN15.05/93393); C, unicuspid rachidian teeth and undifferentiated first lateral of Tritoniopsis elegans (MNHNC/MB28-004506); D, tricuspid rachidian teeth and differentiated first lateral of Tritonidoxa capensis (MNCN15.05/93399); E, unicuspid rachidian teeth and undifferentiated first lateral of Tochuina gigantea (NHMD-115967); F, tricuspid rachidian teeth and differentiated first lateral with several accessory denticles (circled) of Tritonicula hamnerorum (MNCN15.05/93378); G, tricuspid rachidian teeth and differentiated first lateral of Marionia blainvillea (NHMD115968); Scale bars: A, D, E = 60 μm; B, C = 22 μm; F = 20 μm; G = 110 μm." figureDoi="http://doi.org/10.5281/zenodo.8432401" httpUri="https://zenodo.org/record/8432401/files/figure.png" pageId="23" pageNumber="468">4G</figureCitation>
,
<figureCitation id="13342A1FFF876500FD9FFB7AEF88FA93" box="[461,495,1278,1304]" captionStart="Figure 5" captionStartId="14.[129,194,1262,1286]" captionTargetBox="[164,1437,147,1230]" captionTargetId="figure-262@14.[161,1441,144,1234]" captionTargetPageId="14" captionText="Figure 5. Digestive system arrangements of analysed key species of Tritoniinae and Marioniinae subfam. nov., dorsal view: A, Tritonia hombergii (ZMBN 83471); B, Candiella plebeia (MNCN15.05/93393); C, Tritoniopsis flydis (GR7-DMC00040); D, Tritonidoxa capensis(MNCN15.05/93399); E, Tochuina gigantea (NHMD-115967); F, Tritonicula hamnerorum (MNCN15.05/93378); G, Marionia blainvillea (NHMD115968). Abbreviations:a, anus; aodg, anterior ODG complex; e, esophagus; i, intestine; odg, ovotestis-digestive gland complex; s, stomach; podg, posterior ODG complex." figureDoi="http://doi.org/10.5281/zenodo.8432405" httpUri="https://zenodo.org/record/8432405/files/figure.png" pageId="23" pageNumber="468">5G</figureCitation>
,
<figureCitation id="13342A1FFF876500FDAAFB7AEC7EFA93" box="[504,537,1278,1304]" captionStart="Figure 6" captionStartId="15.[113,178,1771,1795]" captionTargetBox="[317,1256,144,1743]" captionTargetId="figure-8@15.[316,1256,144,1743]" captionTargetPageId="15" captionText="Figure 6. Reproductive system arrangements of analysed key species of Tritoniinae and Marioniinae subfam. nov., dorsal view:A, Tritonia hombergii (ZMBN 83471); B, Candiella plebeia (ZMBN 83471); C, Tritoniopsis flydis (GR7-DMC00040); D, Tritonidoxa capensis (MNCN15.05/93399); E, Tochuina gigantea (NHMD-115967); F, Tritonicula hamnerorum (MNCN15.05/93378); G, Marionia blainvillea (NHMD115968). Abbreviations: ag, albumen gland; am, ampulla; bc, bursa copulatrix; fo, female opening; fgm, female gland mass; hd, hermaphrodite duct; o, oviduct; p, penis; pp, prostatic portion; ps, penial sheath; v, vagina; vd, vas deferens." figureDoi="http://doi.org/10.5281/zenodo.8432407" httpUri="https://zenodo.org/record/8432407/files/figure.png" pageId="23" pageNumber="468">6G</figureCitation>
)
</emphasis>
</paragraph>
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<paragraph id="8BB0369AFF876500FC23FAA1EC50FAB6" blockId="23.[113,567,1317,1341]" box="[113,567,1317,1341]" pageId="23" pageNumber="468">
<taxonomicName id="4C0F4D19FF876500FC23FAA1EF30FAB6" ID-CoL="7P387" authority="Risso 1826" authorityName="Risso" authorityYear="1826" box="[113,343,1317,1341]" class="Gastropoda" family="Tritoniidae" genus="Duvaucelia" kingdom="Animalia" order="Nudibranchia" pageId="23" pageNumber="468" phylum="Mollusca" rank="genus">
<emphasis id="B97BEA88FF876500FC23FAA1EEB9FAB6" box="[113,222,1317,1341]" italics="true" pageId="23" pageNumber="468">Duvaucelia</emphasis>
Risso 1826
</taxonomicName>
, Paris, Levrault, 4: 38.
</paragraph>
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<subSubSection id="C3156511FF876500FC23FADBEC9CFA1E" pageId="23" pageNumber="468" type="type_taxon">
<paragraph id="8BB0369AFF876500FC23FADBEC9CFA1E" blockId="23.[113,763,1374,1429]" pageId="23" pageNumber="468">
<emphasis id="B97BEA88FF876500FC23FADBEE8AFAFD" box="[113,237,1375,1398]" italics="true" pageId="23" pageNumber="468">
<typeStatus id="54B48838FF876500FC23FADBEEC6FAFD" box="[113,161,1375,1398]" pageId="23" pageNumber="468">Type</typeStatus>
species:
</emphasis>
<taxonomicName id="4C0F4D19FF876500FCA1FADBEC56FAFD" ID-CoL="735N6" authority="Risso 1818" authorityName="Risso" authorityYear="1818" box="[243,561,1374,1398]" class="Gastropoda" family="Tritoniidae" genus="Marionia" kingdom="Animalia" order="Nudibranchia" pageId="23" pageNumber="468" phylum="Mollusca" rank="species" species="blainvillea">
<emphasis id="B97BEA88FF876500FCA1FADBEFDDFAFD" box="[243,442,1374,1398]" italics="true" pageId="23" pageNumber="468">Marionia blainvillea</emphasis>
Risso 1818
</taxonomicName>
(=
<taxonomicName id="4C0F4D19FF876500FE04FADBEF6BFA1E" ID-CoL="3Y6T9" authority="Vayssiere 1879" authorityName="Vayssiere" authorityYear="1879" class="Gastropoda" family="Tritoniidae" genus="Marionia" kingdom="Animalia" order="Nudibranchia" pageId="23" pageNumber="468" phylum="Mollusca" rank="species" species="berghii">
<emphasis id="B97BEA88FF876500FE04FADBEC9DFAFD" box="[598,762,1374,1398]" italics="true" pageId="23" pageNumber="468">Marionia berghii</emphasis>
<bibRefCitation id="EF9E4B6BFF876500FC23FAF9EF6BFA1E" author="Vayssiere A" box="[113,268,1405,1429]" pageId="23" pageNumber="468" pagination="106 - 118" refId="ref27881" refString="Vayssiere A. Description du Marionia berghii. Journal de Conchyliologie 1879; 27: 106 - 118." type="journal article" year="1879">Vayssière 1879</bibRefCitation>
</taxonomicName>
), by subsequent designation by
<bibRefCitation id="EF9E4B6BFF876500FE1BFAF9EC92FA1E" author="Vayssiere A" box="[585,757,1405,1429]" pageId="23" pageNumber="468" pagination="106 - 118" refId="ref27881" refString="Vayssiere A. Description du Marionia berghii. Journal de Conchyliologie 1879; 27: 106 - 118." type="journal article" year="1879">Vayssière (1879)</bibRefCitation>
.
</paragraph>
</subSubSection>
<subSubSection id="C3156511FF876500FC23FA33ECBCF986" pageId="23" pageNumber="468" type="diagnosis">
<paragraph id="8BB0369AFF876500FC23FA33ECBCF986" blockId="23.[113,763,1463,1550]" pageId="23" pageNumber="468">
<emphasis id="B97BEA88FF876500FC23FA33EEB0FA45" box="[113,215,1463,1486]" italics="true" pageId="23" pageNumber="468">Diagnosis:</emphasis>
Rachidian tooth tricuspid, smooth. Masticatory border denticulate, denticles sharp, triangular or rounded. Digestive system arrangement cladohepatic, U- or G-shaped.
</paragraph>
</subSubSection>
<subSubSection id="C3156511FF876500FC23F9ABEA36FEAE" pageId="23" pageNumber="468" type="description">
<paragraph id="8BB0369AFF876500FC23F9ABECC2F8CA" blockId="23.[113,763,1583,1857]" pageId="23" pageNumber="468">
<emphasis id="B97BEA88FF876500FC23F9ABEE97F9CC" box="[113,240,1583,1607]" italics="true" pageId="23" pageNumber="468">Morphology:</emphasis>
Body slender or broad, length up to
<quantity id="4CF79B7FFF876500FE3CF9ABECA3F9CD" box="[622,708,1583,1607]" metricMagnitude="-1" metricUnit="m" metricValue="2.0" pageId="23" pageNumber="468" unit="mm" value="200.0">200 mm</quantity>
(
<figureCitation id="13342A1FFF876500FE86F9ABEEE0F9ED" captionStart="Figure 3" captionStartId="12.[129,194,1533,1557]" captionTargetBox="[321,1281,144,1505]" captionTargetId="figure-116@12.[321,1281,144,1505]" captionTargetPageId="12" captionText="Figure 3. Representative species of genera of Tritoniidae (with exception of Paratritonia). A, Tritonia hombergii (MNCN15.05/93379) (Portaferry, Ireland); B, Candiella plebeia (MNCN15.05/93393) (Portaferry, Ireland); C, Tritoniopsis elegans (MB28-0045061) (Zavora, Mozambique); D, Tritonidoxa capensis (MNCN15.05/93399) (Scoưburg, South Africa); E, Tritoniella belli (Antarctica); F, Marianina rosea (French Polynesia); G, Myrella challengeriana comb.nov.(Bahía Camarones, Argentina); H, Tochuina gigantea (Agnew Pass, USA); I, Tritonicula khaleesi (MZUSP113165) (Fortaleza, Brazil); J, Marionia cucullata (Golfo Nuevo, Argentina); (K) Marioniopsis levis 1 (MNHNC/ MB28-004895) (Nacala, Mozambique). Photo A and B, courtesy of B. Picton; photo C and K courtesy of Y.Tibiriça; photo D courtesy of V. Fraser; photo E courtesy of M. Caballer; photo F courtesy of G. Rouse; photo G and J, courtesy of N. Baưini; photo H courtesy of S. Thiebaud; photo I by F. Silva." figureDoi="http://doi.org/10.5281/zenodo.8432399" httpUri="https://zenodo.org/record/8432399/files/figure.png" pageId="23" pageNumber="468">Fig. 3J</figureCitation>
). Oral veil bilobed. Four to 18 simple or branched velar processes, with the outermost ones oħen longer. Five to 16 pairs of arborescent gills. Rachidian tooth tricuspid and smooth (
<figureCitation id="13342A1FFF876500FE86F909EEF6F94F" captionStart="Figure 4" captionStartId="13.[113,178,1528,1552]" captionTargetBox="[309,1263,147,1498]" captionTargetId="figure-134@13.[306,1266,144,1501]" captionTargetPageId="13" captionText="Figure 4. Scanning electron micrographs of radulae of analysed key species of Tritoniinae and Marioniinae subfam. nov.: A, tricuspid rachidian teeth and differentiated first lateral of Tritonia hombergii (ZMBN 83471); B, tricuspid rachidian teeth and differentiated first lateral with single accessory denticle (circled) of Candiella plebeia (MNCN15.05/93393); C, unicuspid rachidian teeth and undifferentiated first lateral of Tritoniopsis elegans (MNHNC/MB28-004506); D, tricuspid rachidian teeth and differentiated first lateral of Tritonidoxa capensis (MNCN15.05/93399); E, unicuspid rachidian teeth and undifferentiated first lateral of Tochuina gigantea (NHMD-115967); F, tricuspid rachidian teeth and differentiated first lateral with several accessory denticles (circled) of Tritonicula hamnerorum (MNCN15.05/93378); G, tricuspid rachidian teeth and differentiated first lateral of Marionia blainvillea (NHMD115968); Scale bars: A, D, E = 60 μm; B, C = 22 μm; F = 20 μm; G = 110 μm." figureDoi="http://doi.org/10.5281/zenodo.8432401" httpUri="https://zenodo.org/record/8432401/files/figure.png" pageId="23" pageNumber="468">Fig. 4G</figureCitation>
). Masticatory borders of the jaws denticulate, with medium-sized (up to 50 µm) denticles with sharp cusps and conical or triangular bases. ODG complex cladohepatic, with the stomach located in the upper region of the posterior ODG complex (
<figureCitation id="13342A1FFF876500FE86F88EEEF6F8CA" captionStart="Figure 5" captionStartId="14.[129,194,1262,1286]" captionTargetBox="[164,1437,147,1230]" captionTargetId="figure-262@14.[161,1441,144,1234]" captionTargetPageId="14" captionText="Figure 5. Digestive system arrangements of analysed key species of Tritoniinae and Marioniinae subfam. nov., dorsal view: A, Tritonia hombergii (ZMBN 83471); B, Candiella plebeia (MNCN15.05/93393); C, Tritoniopsis flydis (GR7-DMC00040); D, Tritonidoxa capensis(MNCN15.05/93399); E, Tochuina gigantea (NHMD-115967); F, Tritonicula hamnerorum (MNCN15.05/93378); G, Marionia blainvillea (NHMD115968). Abbreviations:a, anus; aodg, anterior ODG complex; e, esophagus; i, intestine; odg, ovotestis-digestive gland complex; s, stomach; podg, posterior ODG complex." figureDoi="http://doi.org/10.5281/zenodo.8432405" httpUri="https://zenodo.org/record/8432405/files/figure.png" pageId="23" pageNumber="468">Fig. 5G</figureCitation>
). Penis flagelliform (
<figureCitation id="13342A1FFF876500FD3AF8ADEFD5F8CA" box="[360,434,1833,1857]" captionStart="Figure 6" captionStartId="15.[113,178,1771,1795]" captionTargetBox="[317,1256,144,1743]" captionTargetId="figure-8@15.[316,1256,144,1743]" captionTargetPageId="15" captionText="Figure 6. Reproductive system arrangements of analysed key species of Tritoniinae and Marioniinae subfam. nov., dorsal view:A, Tritonia hombergii (ZMBN 83471); B, Candiella plebeia (ZMBN 83471); C, Tritoniopsis flydis (GR7-DMC00040); D, Tritonidoxa capensis (MNCN15.05/93399); E, Tochuina gigantea (NHMD-115967); F, Tritonicula hamnerorum (MNCN15.05/93378); G, Marionia blainvillea (NHMD115968). Abbreviations: ag, albumen gland; am, ampulla; bc, bursa copulatrix; fo, female opening; fgm, female gland mass; hd, hermaphrodite duct; o, oviduct; p, penis; pp, prostatic portion; ps, penial sheath; v, vagina; vd, vas deferens." figureDoi="http://doi.org/10.5281/zenodo.8432407" httpUri="https://zenodo.org/record/8432407/files/figure.png" pageId="23" pageNumber="468">Fig. 6G</figureCitation>
), cylindrical or conical.
</paragraph>
<paragraph id="8BB0369AFF876500FC23F8E7EA36FEAE" blockId="23.[113,763,1891,1977]" lastBlockId="23.[810,1460,144,293]" pageId="23" pageNumber="468">
<emphasis id="B97BEA88FF876500FC23F8E7EC4FF8F0" box="[113,552,1891,1915]" italics="true" pageId="23" pageNumber="468">
Species composition:
<taxonomicName id="4C0F4D19FF876500FD69F8E7EC4FF8F0" box="[315,552,1891,1915]" class="Gastropoda" family="Tritoniidae" genus="Marionia" kingdom="Animalia" order="Nudibranchia" pageId="23" pageNumber="468" phylum="Mollusca" rank="species" species="abrahamorum">Marionia abrahamorum</taxonomicName>
</emphasis>
,
<taxonomicName id="4C0F4D19FF876500FE62F8E7EC92F8F0" authorityName="Risso" authorityYear="1818" box="[560,757,1891,1915]" class="Gastropoda" family="Tritoniidae" genus="Marionia" kingdom="Animalia" order="Nudibranchia" pageId="23" pageNumber="468" phylum="Mollusca" rank="species" species="blainvillea">
<emphasis id="B97BEA88FF876500FE62F8E7EC92F8F0" box="[560,757,1891,1915]" italics="true" pageId="23" pageNumber="468">Marionia blainvillea</emphasis>
</taxonomicName>
,
<taxonomicName id="4C0F4D19FF876500FC23F807EFB9F811" authority="Couthouy 1852" authorityName="Couthouy" authorityYear="1852" box="[113,478,1922,1946]" class="Gastropoda" family="Tritoniidae" genus="Marionia" kingdom="Animalia" order="Nudibranchia" pageId="23" pageNumber="468" phylum="Mollusca" rank="species" species="cucullata">
<emphasis id="B97BEA88FF876500FC23F807EF4AF811" box="[113,301,1922,1946]" italics="true" pageId="23" pageNumber="468">Marionia cucullata</emphasis>
Couthouy 1852
</taxonomicName>
,
<taxonomicName id="4C0F4D19FF876500FDB8F807EEC0F832" authority="Bergh 1902" authorityName="Bergh" authorityYear="1902" class="Gastropoda" family="Tritoniidae" genus="Marionia" kingdom="Animalia" order="Nudibranchia" pageId="23" pageNumber="468" phylum="Mollusca" rank="species" species="chloanthes">
<emphasis id="B97BEA88FF876500FDB8F807ECD2F811" box="[490,693,1922,1946]" italics="true" pageId="23" pageNumber="468">Marionia chloanthes</emphasis>
Bergh 1902
</taxonomicName>
,
<taxonomicName id="4C0F4D19FF876500FCE0F826EC57F832" authority="Jensen 1994" authorityName="Jensen" authorityYear="1994" box="[178,560,1953,1977]" class="Gastropoda" family="Tritoniidae" genus="Marionia" kingdom="Animalia" order="Nudibranchia" pageId="23" pageNumber="468" phylum="Mollusca" rank="species" species="echinomuriceae">
<emphasis id="B97BEA88FF876500FCE0F826EFCEF832" box="[178,425,1953,1977]" italics="true" pageId="23" pageNumber="468">Marionia echinomuriceae</emphasis>
Jensen 1994
</taxonomicName>
,
<taxonomicName id="4C0F4D19FF876500FE68F826EA3BFF23" authority="Edmunds and Carmona 2017" authorityName="Edmunds and Carmona" authorityYear="2017" box="[570,1116,144,1977]" class="Gastropoda" family="Tritoniidae" genus="Marionia" kingdom="Animalia" order="Nudibranchia" pageId="23" pageNumber="468" phylum="Mollusca" rank="species" species="ghanensis">
<emphasis id="B97BEA88FF876500FE68F826EC9DF832" box="[570,762,1953,1977]" italics="true" pageId="23" pageNumber="468">Marionia ghanensis</emphasis>
<bibRefCitation id="EF9E4B6BFF876500FF78FF14EA3BFF23" author="Edmunds M &amp; Carmona L" box="[810,1116,144,168]" pageId="23" pageNumber="468" pagination="339 - 356" refId="ref24940" refString="Edmunds M, Carmona L. Heterobranchia Mollusca from Ghana: the traditionally called Dendronotacea and Arminacea. Journal of Conchology 2017; 42: 339 - 356." type="journal article" year="2017">Edmunds and Carmona 2017</bibRefCitation>
</taxonomicName>
,
<taxonomicName id="4C0F4D19FF876500F834FF15EB68FF23" authorityName="Almon, Perez &amp; Caballer" authorityYear="2018" box="[1126,1295,145,168]" class="Gastropoda" family="Tritoniidae" genus="Marionia" kingdom="Animalia" order="Nudibranchia" pageId="23" pageNumber="468" phylum="Mollusca" rank="species" species="gemmii">
<emphasis id="B97BEA88FF876500F834FF15EB68FF23" box="[1126,1295,145,168]" italics="true" pageId="23" pageNumber="468">Marionia gemmii</emphasis>
</taxonomicName>
,
<taxonomicName id="4C0F4D19FF876500F948FF15EBC9FF23" authorityName="Angulo-Campillo and Bertsch" authorityYear="2013" box="[1306,1454,144,168]" class="Gastropoda" family="Tritoniidae" genus="Marionia" kingdom="Animalia" order="Nudibranchia" pageId="23" pageNumber="468" phylum="Mollusca" rank="species" species="kinoi">
<emphasis id="B97BEA88FF876500F948FF15EBC9FF23" box="[1306,1454,144,168]" italics="true" pageId="23" pageNumber="468">Marionia kinoi</emphasis>
</taxonomicName>
,
<taxonomicName id="4C0F4D19FF876500FF78FF34ED84FF4C" box="[810,995,175,199]" class="Gastropoda" family="Tritoniidae" genus="Marionia" kingdom="Animalia" order="Nudibranchia" pageId="23" pageNumber="468" phylum="Mollusca" rank="species" species="limceana">
<emphasis id="B97BEA88FF876500FF78FF34ED84FF4C" box="[810,995,175,199]" italics="true" pageId="23" pageNumber="468">Marionia limceana</emphasis>
</taxonomicName>
,
<taxonomicName id="4C0F4D19FF876500FFBCFF34EB7EFF4C" authority="Eliot 1904" authorityName="Eliot" authorityYear="1904" box="[1006,1305,175,199]" class="Gastropoda" family="Tritoniidae" genus="Marionia" kingdom="Animalia" order="Nudibranchia" pageId="23" pageNumber="468" phylum="Mollusca" rank="species" species="pellucida">
<emphasis id="B97BEA88FF876500FFBCFF34EAC0FF4C" box="[1006,1191,175,199]" italics="true" pageId="23" pageNumber="468">Marionia pellucida</emphasis>
<bibRefCitation id="EF9E4B6BFF876500F8FEFF2BEB7EFF4C" author="Eliot C" box="[1196,1305,175,199]" pageId="23" pageNumber="468" pagination="268 - 298" refId="ref24969" refString="Eliot C. On some nudibranchs from east Africa and Zanzibar, Part VI. Proceedings of the Zoological Society of London 1904; 74: 268 - 298." type="journal article" year="1904">Eliot 1904</bibRefCitation>
</taxonomicName>
,
<taxonomicName id="4C0F4D19FF876500F970FF34EB19FF4C" authorityName="Vayssiere" authorityYear="1877" box="[1314,1406,176,199]" class="Gastropoda" family="Tritoniidae" genus="Marionia" kingdom="Animalia" order="Nudibranchia" pageId="23" pageNumber="468" phylum="Mollusca" rank="genus">
<emphasis id="B97BEA88FF876500F970FF34EB19FF4C" box="[1314,1406,176,199]" italics="true" pageId="23" pageNumber="468">Marionia</emphasis>
</taxonomicName>
<taxonomicName id="4C0F4D19FF876500F9D6FF34EA08FF6D" authority="Er. Marcus and Ev. Marcus 1968" authorityName="Er. Marcus and Ev. Marcus" authorityYear="1968" class="Gastropoda" family="Tritoniidae" genus="Marionia" kingdom="Animalia" order="Nudibranchia" pageId="23" pageNumber="468" phylum="Mollusca" rank="species" species="pusa">
pusa Er. Marcus and Ev.
<bibRefCitation id="EF9E4B6BFF876500FFBAFF4BEA08FF6D" author="Marcus Ev &amp; Marcus Er" box="[1000,1135,206,230]" pageId="23" pageNumber="468" pagination="1334 - 1342" refId="ref26080" refString="Marcus Ev, Marcus Er. Some opisthobranchs from Ivory Coast. Bulletin de l'Institut Fondamental de l'Aflique Noire (A) 1968; 30: 1334 - 1342." type="journal article" year="1968">Marcus 1968</bibRefCitation>
</taxonomicName>
,
<taxonomicName id="4C0F4D19FF876500F825FF4BEDA8FE8D" authority="Ev. Marcus and Er. Marcus 1966" authorityName="Ev. Marcus and Er. Marcus" authorityYear="1966" class="Gastropoda" family="Tritoniidae" genus="Marionia" kingdom="Animalia" order="Nudibranchia" pageId="23" pageNumber="468" phylum="Mollusca" rank="species" species="vanira">
<emphasis id="B97BEA88FF876500F825FF4BEB72FF6D" box="[1143,1301,207,230]" italics="true" pageId="23" pageNumber="468">Marionia vanira</emphasis>
Ev. Marcus and Er.
<bibRefCitation id="EF9E4B6BFF876500FF19FF6AEDA8FE8D" author="Marcus Ev &amp; Marcus Er" box="[843,975,238,262]" pageId="23" pageNumber="468" pagination="152 - 208" refId="ref26018" refString="Marcus Ev, Marcus Er. Opisthobranchs from tropical West Africa. Studies in Tropical Oceanography 1966; 4: 152 - 208." type="journal article" year="1966">Marcus 1966</bibRefCitation>
</taxonomicName>
and the new species
<taxonomicName id="4C0F4D19FF876500F8CDFF6BEBF3FE8D" authority="Silva et al." authorityName="Silva" authorityYear="2023" box="[1183,1428,238,262]" class="Gastropoda" family="Tritoniidae" genus="Marionia" kingdom="Animalia" order="Nudibranchia" pageId="23" pageNumber="468" phylum="Mollusca" rank="species" species="flani" status="sp. nov.">
<emphasis id="B97BEA88FF876500F8CDFF6BEB49FE8D" box="[1183,1326,238,262]" italics="true" pageId="23" pageNumber="468">Marionia flani</emphasis>
Silva
<emphasis id="B97BEA88FF876500F935FF6BEBF3FE8D" box="[1383,1428,238,262]" italics="true" pageId="23" pageNumber="468">et al.</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A24857F3FF876500F9CAFF6AED32FEAE" pageId="23" pageNumber="468" rank="species">sp. nov.</taxonomicNameLabel>
(see description below).
</paragraph>
</subSubSection>
<subSubSection id="C3156511FF876500FF78FEC3EBC8FB06" pageId="23" pageNumber="468" type="discussion">
<paragraph id="8BB0369AFF876500FF78FEC3EB5FFD90" blockId="23.[810,1461,327,1165]" pageId="23" pageNumber="468">
<emphasis id="B97BEA88FF876500FF78FEC3EDEFFED4" box="[810,904,327,351]" italics="true" pageId="23" pageNumber="468">Remarks:</emphasis>
The morphological synapomorphies of
<taxonomicName id="4C0F4D19FF876500F905FEC3EBD4FED5" authorityName="Vayssiere" authorityYear="1877" box="[1367,1459,327,350]" class="Gastropoda" family="Tritoniidae" genus="Marionia" kingdom="Animalia" order="Nudibranchia" pageId="23" pageNumber="468" phylum="Mollusca" rank="genus">
<emphasis id="B97BEA88FF876500F905FEC3EBD4FED5" box="[1367,1459,327,350]" italics="true" pageId="23" pageNumber="468">Marionia</emphasis>
</taxonomicName>
within
<taxonomicName id="4C0F4D19FF876500FF25FEE2ED92FEF5" box="[887,1013,358,382]" class="Gastropoda" family="Tritoniidae" kingdom="Animalia" order="Nudibranchia" pageId="23" pageNumber="467" phylum="Mollusca" rank="subFamily" subFamily="Marioniinae">Marioniinae</taxonomicName>
are the smooth tricuspid rachidian tooth (
<figureCitation id="13342A1FFF876500FF67FE01EDE7FE16" box="[821,896,389,413]" captionStart="Figure 4" captionStartId="13.[113,178,1528,1552]" captionTargetBox="[309,1263,147,1498]" captionTargetId="figure-134@13.[306,1266,144,1501]" captionTargetPageId="13" captionText="Figure 4. Scanning electron micrographs of radulae of analysed key species of Tritoniinae and Marioniinae subfam. nov.: A, tricuspid rachidian teeth and differentiated first lateral of Tritonia hombergii (ZMBN 83471); B, tricuspid rachidian teeth and differentiated first lateral with single accessory denticle (circled) of Candiella plebeia (MNCN15.05/93393); C, unicuspid rachidian teeth and undifferentiated first lateral of Tritoniopsis elegans (MNHNC/MB28-004506); D, tricuspid rachidian teeth and differentiated first lateral of Tritonidoxa capensis (MNCN15.05/93399); E, unicuspid rachidian teeth and undifferentiated first lateral of Tochuina gigantea (NHMD-115967); F, tricuspid rachidian teeth and differentiated first lateral with several accessory denticles (circled) of Tritonicula hamnerorum (MNCN15.05/93378); G, tricuspid rachidian teeth and differentiated first lateral of Marionia blainvillea (NHMD115968); Scale bars: A, D, E = 60 μm; B, C = 22 μm; F = 20 μm; G = 110 μm." figureDoi="http://doi.org/10.5281/zenodo.8432401" httpUri="https://zenodo.org/record/8432401/files/figure.png" pageId="23" pageNumber="468">Fig. 4G</figureCitation>
), sharp and triangular or rounded denticles in the masticatory border and a unique arrangement of the cladohepatic digestive system: the stomach and intestine are located above the upper region of the posterior ODG complex, and the entire arrangement is in the shape of a U or G (
<figureCitation id="13342A1FFF876500F88FFD86EB4FFD91" box="[1245,1320,514,538]" captionStart="Figure 5" captionStartId="14.[129,194,1262,1286]" captionTargetBox="[164,1437,147,1230]" captionTargetId="figure-262@14.[161,1441,144,1234]" captionTargetPageId="14" captionText="Figure 5. Digestive system arrangements of analysed key species of Tritoniinae and Marioniinae subfam. nov., dorsal view: A, Tritonia hombergii (ZMBN 83471); B, Candiella plebeia (MNCN15.05/93393); C, Tritoniopsis flydis (GR7-DMC00040); D, Tritonidoxa capensis(MNCN15.05/93399); E, Tochuina gigantea (NHMD-115967); F, Tritonicula hamnerorum (MNCN15.05/93378); G, Marionia blainvillea (NHMD115968). Abbreviations:a, anus; aodg, anterior ODG complex; e, esophagus; i, intestine; odg, ovotestis-digestive gland complex; s, stomach; podg, posterior ODG complex." figureDoi="http://doi.org/10.5281/zenodo.8432405" httpUri="https://zenodo.org/record/8432405/files/figure.png" pageId="23" pageNumber="468">Fig. 5G</figureCitation>
).
</paragraph>
<paragraph id="8BB0369AFF876500FF14FDA6EBC8FB06" blockId="23.[810,1461,327,1165]" pageId="23" pageNumber="468">
According to our phylogenetic analyses,
<taxonomicName id="4C0F4D19FF876500F8B7FDA7EB26FDB1" authorityName="Vayssiere" authorityYear="1877" box="[1253,1345,547,570]" class="Gastropoda" family="Tritoniidae" genus="Marionia" kingdom="Animalia" order="Nudibranchia" pageId="23" pageNumber="468" phylum="Mollusca" rank="genus">
<emphasis id="B97BEA88FF876500F8B7FDA7EB26FDB1" box="[1253,1345,547,570]" italics="true" pageId="23" pageNumber="468">Marionia</emphasis>
</taxonomicName>
consists of two clades with different geographical compositions: one clade includes the Pacific
<taxonomicName id="4C0F4D19FF876500FFAAFDE5EAEBFDF3" authorityName="Angulo-Campillo and Bertsch" authorityYear="2013" box="[1016,1164,608,632]" class="Gastropoda" family="Tritoniidae" genus="Marionia" kingdom="Animalia" order="Nudibranchia" pageId="23" pageNumber="468" phylum="Mollusca" rank="species" species="kinoi">
<emphasis id="B97BEA88FF876500FFAAFDE5EAEBFDF3" box="[1016,1164,608,632]" italics="true" pageId="23" pageNumber="468">Marionia kinoi</emphasis>
</taxonomicName>
and several Atlantic species, including the
<typeStatus id="54B48838FF876500FF96FD04ED95FD13" box="[964,1010,640,664]" pageId="23" pageNumber="468">type</typeStatus>
species
<taxonomicName id="4C0F4D19FF876500F801FD05EB46FD13" authorityName="Risso" authorityYear="1818" box="[1107,1313,640,664]" class="Gastropoda" family="Tritoniidae" genus="Marionia" kingdom="Animalia" order="Nudibranchia" pageId="23" pageNumber="468" phylum="Mollusca" rank="species" species="blainvillea">
<emphasis id="B97BEA88FF876500F801FD05EB46FD13" box="[1107,1313,640,664]" italics="true" pageId="23" pageNumber="468">Marionia blainvillea</emphasis>
</taxonomicName>
, whereas the other is comprised of the Atlantic
<taxonomicName id="4C0F4D19FF876500F8DFFD24EB20FD3C" box="[1165,1351,671,695]" class="Gastropoda" family="Tritoniidae" genus="Marionia" kingdom="Animalia" order="Nudibranchia" pageId="23" pageNumber="468" phylum="Mollusca" rank="species" species="limceana">
<emphasis id="B97BEA88FF876500F8DFFD24EB20FD3C" box="[1165,1351,671,695]" italics="true" pageId="23" pageNumber="468">Marionia limceana</emphasis>
</taxonomicName>
and many unnamed Pacific species (
<figureCitation id="13342A1FFF876500F810FD3AEAE7FD5C" box="[1090,1152,702,727]" captionStart="Figure 2" captionStartId="10.[129,194,1719,1743]" captionTargetBox="[325,1278,148,1689]" captionTargetId="figure-6@10.[321,1281,144,1692]" captionTargetPageId="10" captionText="Figure 2. Bayesian phylogenetic tree based on the concatenated molecular data (COI + 16S + H3) of Tritoniidae (sensu Odhner 1963). Outgroup taxa are depicted in grey.Tritoniidae genera are independently coloured, and branches with several replicates for each species hypothesis were collapsed for clarity.Names in bold are species which sequences were generated for this study with the number of sequenced specimens showed in brackets. PP are shown right/top and BS leħ/boưom.Unsupported branches are not labelled. Green circles indicate maximum support by both analyses. Solid vertical bars indicate specimens grouped as species entities based on COI dataset. Empty vertical bars indicate the specimens grouped as species based on the 16S dataset." figureDoi="http://doi.org/10.5281/zenodo.8432395" httpUri="https://zenodo.org/record/8432395/files/figure.png" pageId="23" pageNumber="468">Fig. 2</figureCitation>
). The inclusion of
<taxonomicName id="4C0F4D19FF876500F905FD3BEDE6FD7D" authorityName="Couthouy" authorityYear="1852" class="Gastropoda" family="Tritoniidae" genus="Marionia" kingdom="Animalia" order="Nudibranchia" pageId="23" pageNumber="468" phylum="Mollusca" rank="species" species="cucullata">
<emphasis id="B97BEA88FF876500F905FD3BEDE6FD7D" italics="true" pageId="23" pageNumber="468">Marionia cucullata</emphasis>
</taxonomicName>
in the genus is based on a recent phylogenetic hypothesis that shows the species grouped with
<taxonomicName id="4C0F4D19FF876500F8BAFD7AED95FCBE" authority="(Moles et al. 2021)" baseAuthorityName="Moles" baseAuthorityYear="2021" class="Gastropoda" family="Tritoniidae" genus="Marionia" kingdom="Animalia" order="Nudibranchia" pageId="23" pageNumber="468" phylum="Mollusca" rank="species" species="blainvillea">
<emphasis id="B97BEA88FF876500F8BAFD7AEBD4FC9E" box="[1256,1459,765,789]" italics="true" pageId="23" pageNumber="468">Marionia blainvillea</emphasis>
(
<bibRefCitation id="EF9E4B6BFF876500FF67FC98ED80FCBE" author="Moles J &amp; Berning MI &amp; Hooker Y" box="[821,999,796,821]" pageId="23" pageNumber="468" pagination="107209" refId="ref26342" refString="Moles J, Berning MI, Hooker Y, et al. Due South: the evolutionary history of sub-Antarctic and Antarctic Tritoniidae nudibranchs. Molecular Phylogenetics and Evolution 2021; 162: 107209." type="journal article" year="2021">
Moles
<emphasis id="B97BEA88FF876500FF2EFC99EDC1FCBF" box="[892,934,796,820]" italics="true" pageId="23" pageNumber="468">et al</emphasis>
. 2021
</bibRefCitation>
)
</taxonomicName>
, as well as on morphological evidence that fits our proposed diagnosis: the G-shaped arrangement of the digestive system and the smooth tricuspid rachidian tooth (
<bibRefCitation id="EF9E4B6BFF876500F933FCDFED39FC18" author="Odhner NH" pageId="23" pageNumber="468" pagination="229 - 310" refId="ref26550" refString="Odhner NH. The Nudibranchiata of British Antarctic Expedition. British Antartic (' Terra Nova') Expedition, 1910. Natural History Reports. Zoology 1934; 7: 229 - 310." type="journal article" year="1934">Odhner 1934</bibRefCitation>
). Species for which molecular data are not available, namely
<taxonomicName id="4C0F4D19FF876500FF2EFC1EEA21FC39" authorityName="Bergh" authorityYear="1902" box="[892,1094,922,946]" class="Gastropoda" family="Tritoniidae" genus="Marionia" kingdom="Animalia" order="Nudibranchia" pageId="23" pageNumber="468" phylum="Mollusca" rank="species" species="chloanthes">
<emphasis id="B97BEA88FF876500FF2EFC1EEA21FC39" box="[892,1094,922,946]" italics="true" pageId="23" pageNumber="468">Marionia chloanthes</emphasis>
</taxonomicName>
,
<taxonomicName id="4C0F4D19FF876500F800FC1EEB2DFC39" authorityName="Jensen" authorityYear="1994" box="[1106,1354,922,946]" class="Gastropoda" family="Tritoniidae" genus="Marionia" kingdom="Animalia" order="Nudibranchia" pageId="23" pageNumber="468" phylum="Mollusca" rank="species" species="echinomuriceae">
<emphasis id="B97BEA88FF876500F800FC1EEB2DFC39" box="[1106,1354,922,946]" italics="true" pageId="23" pageNumber="468">Marionia echinomuriceae</emphasis>
</taxonomicName>
,
<taxonomicName id="4C0F4D19FF876500F905FC1EEDE0FC5A" authorityName="Edmunds and Carmona" authorityYear="2017" class="Gastropoda" family="Tritoniidae" genus="Marionia" kingdom="Animalia" order="Nudibranchia" pageId="23" pageNumber="468" phylum="Mollusca" rank="species" species="ghanensis">
<emphasis id="B97BEA88FF876500F905FC1EEDE0FC5A" italics="true" pageId="23" pageNumber="468">Marionia ghanensis</emphasis>
</taxonomicName>
,
<taxonomicName id="4C0F4D19FF876500FFC3FC3EEA2FFC5A" authorityName="Eliot" authorityYear="1904" box="[913,1096,953,977]" class="Gastropoda" family="Tritoniidae" genus="Marionia" kingdom="Animalia" order="Nudibranchia" pageId="23" pageNumber="468" phylum="Mollusca" rank="species" species="pellucida">
<emphasis id="B97BEA88FF876500FFC3FC3EEA2FFC5A" box="[913,1096,953,977]" italics="true" pageId="23" pageNumber="468">Marionia pellucida</emphasis>
</taxonomicName>
,
<taxonomicName id="4C0F4D19FF876500F800FC3EEA87FC5A" authorityName="Er. Marcus and Ev. Marcus" authorityYear="1968" box="[1106,1248,954,977]" class="Gastropoda" family="Tritoniidae" genus="Marionia" kingdom="Animalia" order="Nudibranchia" pageId="23" pageNumber="468" phylum="Mollusca" rank="species" species="pusa">
<emphasis id="B97BEA88FF876500F800FC3EEA87FC5A" box="[1106,1248,954,977]" italics="true" pageId="23" pageNumber="468">Marionia pusa</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0F4D19FF876500F95DFC3EEBCAFC5A" authorityName="Ev. Marcus and Er. Marcus" authorityYear="1966" box="[1295,1453,954,977]" class="Gastropoda" family="Tritoniidae" genus="Marionia" kingdom="Animalia" order="Nudibranchia" pageId="23" pageNumber="468" phylum="Mollusca" rank="species" species="vanira">
<emphasis id="B97BEA88FF876500F95DFC3EEBCAFC5A" box="[1295,1453,954,977]" italics="true" pageId="23" pageNumber="468">Marionia vanira</emphasis>
</taxonomicName>
, are included in this genus on the basis of the presence of true stomach plates, the cladohepatic ODG complex and the smooth tricuspid rachidian tooth (Ev. Marcus and Er.
<bibRefCitation id="EF9E4B6BFF876500F950FB9CEBEEFBA4" author="Marcus Ev &amp; Marcus Er" box="[1282,1417,1047,1071]" pageId="23" pageNumber="468" pagination="152 - 208" refId="ref26018" refString="Marcus Ev, Marcus Er. Opisthobranchs from tropical West Africa. Studies in Tropical Oceanography 1966; 4: 152 - 208." type="journal article" year="1966">Marcus 1966</bibRefCitation>
; Er. Marcus and Ev.
<bibRefCitation id="EF9E4B6BFF876500FF83FBB3EA3EFBC5" author="Marcus Ev &amp; Marcus Er" box="[977,1113,1078,1102]" pageId="23" pageNumber="468" pagination="1334 - 1342" refId="ref26080" refString="Marcus Ev, Marcus Er. Some opisthobranchs from Ivory Coast. Bulletin de l'Institut Fondamental de l'Aflique Noire (A) 1968; 30: 1334 - 1342." type="journal article" year="1968">Marcus 1968</bibRefCitation>
; Ev.
<bibRefCitation id="EF9E4B6BFF876500F8DFFBB3EB72FBC5" author="Marcus Ev" box="[1165,1301,1078,1102]" pageId="23" pageNumber="468" pagination="177 - 214" refId="ref25994" refString="Marcus Ev. The Western Atlantic Tritoniidae. Boletim de Zoologia, Universidade de Sao Paulo 1983; 6: 177 - 214." type="journal article" year="1983">Marcus 1983</bibRefCitation>
,
<bibRefCitation id="EF9E4B6BFF876500F970FBB2EDA3FBE5" author="Edmunds M &amp; Carmona L" pageId="23" pageNumber="468" pagination="339 - 356" refId="ref24940" refString="Edmunds M, Carmona L. Heterobranchia Mollusca from Ghana: the traditionally called Dendronotacea and Arminacea. Journal of Conchology 2017; 42: 339 - 356." type="journal article" year="2017">Edmunds and Carmona 2017</bibRefCitation>
).
<taxonomicName id="4C0F4D19FF876500FF88FBD2EA1BFBE6" box="[986,1148,1110,1133]" class="Gastropoda" family="Tritoniidae" genus="Marionia" kingdom="Animalia" order="Nudibranchia" pageId="23" pageNumber="468" phylum="Mollusca" rank="species" species="gemmi">
<emphasis id="B97BEA88FF876500FF88FBD2EA1BFBE6" box="[986,1148,1110,1133]" italics="true" pageId="23" pageNumber="468">Marionia gemmi</emphasis>
</taxonomicName>
is the only species of
<taxonomicName id="4C0F4D19FF876500F905FBD2EBD4FBE6" authorityName="Vayssiere" authorityYear="1877" box="[1367,1459,1110,1133]" class="Gastropoda" family="Tritoniidae" genus="Marionia" kingdom="Animalia" order="Nudibranchia" pageId="23" pageNumber="468" phylum="Mollusca" rank="genus">
<emphasis id="B97BEA88FF876500F905FBD2EBD4FBE6" box="[1367,1459,1110,1133]" italics="true" pageId="23" pageNumber="468">Marionia</emphasis>
</taxonomicName>
with smooth masticatory border of the jaw (
<bibRefCitation id="EF9E4B6BFF876500F8B9FBF1EBF9FB06" author="Almon B &amp; Perez J &amp; Caballer M" box="[1259,1438,1141,1165]" pageId="23" pageNumber="468" pagination="892 - 908" refId="ref24068" refString="Almon B, Perez J, Caballer M. Expect the unexpected: a new large species of Marionia (Heterobranchia: Nudibranchia: Tritoniidae) from Western Europe. Invertebrate Systematics 2018; 32: 892 - 908." type="journal article" year="2018">
Almón
<emphasis id="B97BEA88FF876500F965FBF2EB38FB06" box="[1335,1375,1141,1165]" italics="true" pageId="23" pageNumber="468">et al</emphasis>
. 2018
</bibRefCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>