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<mods:title id="9A5B8E6E1EC64DA673B866063B51C400">A synoptic review of the family Dendronotidae (Mollusca: Nudibranchia): a multilevel organismal diversity approach</mods:title>
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<paragraph id="8BBEA6780244FF8F21B1FA21CE6514F9" blockId="13.[645,1139,1499,1526]" box="[645,1139,1499,1526]" pageId="13" pageNumber="106">
<heading id="D0F611140244FF8F21B1FA21CE6514F9" box="[645,1139,1499,1526]" pageId="13" pageNumber="106">
<emphasis id="B9757A6A0244FF8F21B1FA21CE6514F9" bold="true" box="[645,1139,1499,1526]" pageId="13" pageNumber="106">
Genus
<taxonomicName id="4C01DDFB0244FF8F21E0FA26CE6514F9" ID-CoL="7NVQ3" authority="Alder &amp; Hancock, 1845" authorityName="Alder &amp; Hancock" authorityYear="1845" box="[724,1139,1499,1526]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="13" pageNumber="106" phylum="Mollusca" rank="genus">
<emphasis id="B9757A6A0244FF8F21E0FA26C97F14F9" bold="true" box="[724,873,1499,1526]" italics="true" pageId="13" pageNumber="106">Dendronotus</emphasis>
Alder &amp; Hancock, 1845
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<emphasis id="B9757A6A0244FF8F21B1F9FCC91D1715" box="[645,779,1537,1562]" italics="true" pageId="13" pageNumber="106">
<typeStatus id="54BA18DA0244FF8F21B1F9FCC8AF1715" box="[645,697,1537,1562]" pageId="13" pageNumber="106">Type</typeStatus>
species
</emphasis>
.
<taxonomicName id="4C01DDFB0244FF8F2023F9FDC9201731" ID-CoL="34QZD" authority="(Ascanius, 1774)" baseAuthorityName="Ascanius" baseAuthorityYear="1774" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="13" pageNumber="106" phylum="Mollusca" rank="species" species="frondosus">
<emphasis id="B9757A6A0244FF8F2023F9FDCE031715" box="[791,1045,1536,1562]" italics="true" pageId="13" pageNumber="106">Dendronotusfrondosus</emphasis>
(
<bibRefCitation id="EF90DB890244FF8F21B9F9D9C9381731" author="Ascanius, P." box="[653,814,1572,1598]" journalOrPublisher="Det Kongelige Norske Videnskabelige Selskabs Skrifter Trondhejm" pageId="13" pageNumber="106" pagination="153 - 158" part="5" refId="ref27923" refString="Ascanius, P. (1774) Beskrivelse over en Norske sneppe og et sodyr. Det Kongelige Norske Videnskabelige Selskabs Skrifter Trondhejm, 5, 153 - 158." title="Beskrivelse over en Norske sneppe og et sodyr" type="journal article" year="1774">Ascanius, 1774</bibRefCitation>
)
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</paragraph>
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<paragraph id="8BBEA6780244FF8F21B1F991CE9317C2" blockId="13.[645,1157,1643,1742]" pageId="13" pageNumber="106">
<emphasis id="B9757A6A0244FF8F21B1F991C8FA178A" box="[645,748,1644,1669]" italics="true" pageId="13" pageNumber="106">Diagnosis</emphasis>
.Dorsolateralappendageswithdistinct tertiary branches. Oral veil with branched appendages. Jaws commonly with set of tightly
</paragraph>
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<paragraph id="8BBEA6780247FF8C23B5F979C92A17DC" blockId="14.[129,1121,1668,1748]" pageId="14" pageNumber="107">
<pageTitle id="CB9E7E1F0247FF8C23B5F979CAC41796" box="[129,210,1668,1689]" pageId="14" pageNumber="107">FIGURE 7</pageTitle>
Representaives of the genus
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<emphasis id="B9757A6A0247FF8C22C9F979C8671796" box="[509,625,1668,1689]" italics="true" pageId="14" pageNumber="107">Dendronotus</emphasis>
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(living specimens). PHOTOGRAPHS BY T. KORSHUNOVA, A. MARTYNOV, K. FLETCHER, D. MILLER, Y. FUJIWARA, K. HASEGAWA, K. SANAMYAN, N. SANAMYAN, AND O. ZIMINA
</paragraph>
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<paragraph id="8BBEA6780246FF8D2356FF56CBBE120B" blockId="15.[98,611,171,772]" pageId="15" pageNumber="108">packed ridge-like semi-discoid structures and also with denticles. Radula with at least more than eight lateral teeth in adult specimens. Central teethwith cusp integrated withinlateral denticles (or within teeth body if denticles absent), the central cusp is not protruding. Denticle-bear part of central teeth separated from lateral sides by usually distinct shoulder, thus central teeth with squarish base and triangular top.Prostatewith evident, conspicous alveolar glands, commonly represent by a discoid structure (varies from a flattened disk to a compressed spherical structure), or very rarely non-discoid, linear (in this case alveolar glands also present). Copulative organ basically conical, with various ratios of length and width anddegrees of apical tapering.</paragraph>
</subSubSection>
<subSubSection id="C31BF5F30246FF902356FCCFC9011732" lastPageId="18" lastPageNumber="111" pageId="15" pageNumber="108" type="discussion">
<paragraph id="8BBEA6780246FF8D2356FCCFCB671485" blockId="15.[98,611,818,1418]" pageId="15" pageNumber="108">
<emphasis id="B9757A6A0246FF8D2356FCCFCB1D1243" box="[98,267,818,844]" italics="true" pageId="15" pageNumber="108">Natural history</emphasis>
. Representaives of the genus
<taxonomicName id="4C01DDFB0246FF8D2356FCABCAF9127F" authorityName="Alder &amp; Hancock" authorityYear="1845" box="[98,239,854,880]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="15" pageNumber="108" phylum="Mollusca" rank="genus">
<emphasis id="B9757A6A0246FF8D2356FCABCAF9127F" box="[98,239,854,880]" italics="true" pageId="15" pageNumber="108">Dendronotus</emphasis>
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inhabit a broad bathymetric range, from intertidal to depths over
<quantity id="4CF90B9D0246FF8D2132FC87C84B129B" box="[518,605,890,916]" metricMagnitude="3" metricUnit="m" metricValue="3.0" pageId="15" pageNumber="108" unit="m" value="3000.0">3000 m</quantity>
, and a variety of substrates (including stony and soft benthic environments). Amajority of the
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<emphasis id="B9757A6A0246FF8D2385FC18CB2812F0" box="[177,318,997,1023]" italics="true" pageId="15" pageNumber="108">Dendronotus</emphasis>
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species feed on hydroids (
<bibRefCitation id="EF90DB890246FF8D235EFBF4CB0B152C" author="Robilliard, G." box="[106,285,1033,1059]" journalOrPublisher="Veliger" pageId="15" pageNumber="108" pagination="433 - 479" part="12" refId="ref31995" refString="Robilliard, G. (1970) The systematics and some aspects of the ecology of the genus Dendronotus. Veliger, 12, 433 - 479." title="The systematics and some aspects of the ecology of the genus Dendronotus" type="journal article" year="1970">Robilliard, 1970</bibRefCitation>
;
<bibRefCitation id="EF90DB890246FF8D2218FBF4C848152C" author="Thompson, T. &amp; Brown, G." box="[300,606,1033,1060]" journalOrPublisher="The Ray Society Publishing, London" pageId="15" pageNumber="108" refId="ref32859" refString="Thompson, T. &amp; Brown, G. (1984) Biology of Opisthobranch Molluscs (vol. 2). The Ray Society Publishing, London." title="Biology of Opisthobranch Molluscs (vol. 2)" type="book" year="1984">Thompson &amp; Brown, 1984</bibRefCitation>
;
<bibRefCitation id="EF90DB890246FF8D2356FBD3CBA41548" author="Martynov, A. V. &amp; Korshunova, T. A." box="[98,434,1069,1096]" journalOrPublisher="Fiton, Moscow" pageId="15" pageNumber="108" refId="ref30931" refString="Martynov, A. V. &amp; Korshunova, T. A. (2011) Opisthobranch Molluscs of the Seas of Russia. A Colour Guide to their Taxonomy and Biology. Fiton, Moscow." title="Opisthobranch Molluscs of the Seas of Russia. A Colour Guide to their Taxonomy and Biology" type="book" year="2011">Martynov &amp; Korshunova, 2011</bibRefCitation>
). There as a few notable exceptions, such as
<taxonomicName id="4C01DDFB0246FF8D22A1FBAFC8771564" authority="Cooper, 1863" authorityName="Cooper" authorityYear="1863" box="[405,609,1105,1131]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="15" pageNumber="108" phylum="Mollusca" rank="species" species="iris">
<emphasis id="B9757A6A0246FF8D22A1FBAFCBC41564" box="[405,466,1106,1131]" italics="true" pageId="15" pageNumber="108">D. iris</emphasis>
<bibRefCitation id="EF90DB890246FF8D22E3FBACC8771564" author="Cooper, J." box="[471,609,1105,1131]" journalOrPublisher="Proc. Calif. Acad. Nat. Sci." pageId="15" pageNumber="108" pagination="56 - 60" part="3" refId="ref28557" refString="Cooper, J. (1863) On new or rare Mollusca inhabiting the coast of California. No. II. Proc. Calif. Acad. Nat. Sci., 3, 56 - 60." title="On new or rare Mollusca inhabiting the coast of California. No. II" type="journal article" year="1863">Cooper, 1863</bibRefCitation>
</taxonomicName>
fromthenortheasternPacific,whichisanactive predator of ceriantharian cnidarians (
<bibRefCitation id="EF90DB890246FF8D2137FB64CA8215D8" author="Wobber, D." journalOrPublisher="Veliger" pageId="15" pageNumber="108" pagination="383 - 387" part="12" refId="ref33357" refString="Wobber, D. (1970) A report on the feeding of Dendronotus iris on the anthozoan Cerianthus sp. from Monterey Bay, California. Veliger, 12, 383 - 387." title="A report on the feeding of Dendronotus iris on the anthozoan Cerianthus sp. from Monterey Bay, California" type="journal article" year="1970">Wobber, 1970</bibRefCitation>
;
<bibRefCitation id="EF90DB890246FF8D23AAFB40CB0515D8" author="Shaw, G." box="[158,275,1213,1239]" journalOrPublisher="Can. J. Zool." pageId="15" pageNumber="108" pagination="2805 - 2810" part="69" refId="ref32501" refString="Shaw, G. (1991) Chemotaxis and lunge-feeding behaviour of Dendronotus iris (Mollusca, Opisthobranchia). Can. J. Zool., 69, 2805 - 2810." title="Chemotaxis and lunge-feeding behaviour of Dendronotus iris (Mollusca, Opisthobranchia)" type="journal article" year="1991">Shaw, 1991</bibRefCitation>
), whereas for the wide-bodied species
<taxonomicName id="4C01DDFB0246FF8D238EFB1CCBA915F4" authority="Verrill, 1870" authorityName="Verrill" authorityYear="1870" box="[186,447,1248,1275]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="15" pageNumber="108" phylum="Mollusca" rank="species" species="robustus">
<emphasis id="B9757A6A0246FF8D238EFB1CCB2315F5" box="[186,309,1248,1274]" italics="true" pageId="15" pageNumber="108">D. robustus</emphasis>
<bibRefCitation id="EF90DB890246FF8D2209FB1DCBA915F4" author="Verrill, A." box="[317,447,1248,1275]" journalOrPublisher="Descriptions of some New England Nudibranchiata. Am. J. Sci. Arts" pageId="15" pageNumber="108" pagination="405 - 408" part="50" refId="ref33156" refString="Verrill, A. (1870) Contributions to zoology from the Museum of Yale College. No. 8. - Descriptions of some New England Nudibranchiata. Am. J. Sci. Arts (Ser. 2), 50, 405 - 408." title="Contributions to zoology from the Museum of Yale College" type="journal article" year="1870">Verrill, 1870</bibRefCitation>
</taxonomicName>
,
<taxonomicName id="4C01DDFB0246FF8D22FFFB1CCAC61410" authority="G. O. Sars, 1878" authorityName="G. O. Sars" authorityYear="1878" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="15" pageNumber="108" phylum="Mollusca" rank="species" species="velifer">
<emphasis id="B9757A6A0246FF8D22FFFB1CC83C15F5" box="[459,554,1248,1274]" italics="true" pageId="15" pageNumber="108">D. velifer</emphasis>
G.O.
<bibRefCitation id="EF90DB890246FF8D2356FAF8CAC61410" author="Sars, G." box="[98,208,1285,1311]" journalOrPublisher="BlOddyr. Universitetsprogram, Christiania" pageId="15" pageNumber="108" refId="ref32434" refString="Sars, G. (1878) Bidrag til kundskaben om Norges Arktiske fauna. I. Mollusca regionis Arcticae Norvegiae, oversigtoverdeiNorgesArktiskeregion forekommende. BlOddyr. Universitetsprogram, Christiania." title="Bidrag til kundskaben om Norges Arktiske fauna. I. Mollusca regionis Arcticae Norvegiae, oversigtoverdeiNorgesArktiskeregion forekommende" type="book" year="1878">Sars, 1878</bibRefCitation>
</taxonomicName>
and
<taxonomicName id="4C01DDFB0246FF8D2226FAF8CAB2144C" authority="Martynov et al., 2020 a" authorityName="Martynov" authorityYear="2020" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="15" pageNumber="108" phylum="Mollusca" rank="species" species="bathyvela">
<emphasis id="B9757A6A0246FF8D2226FAF8CB8B1411" box="[274,413,1284,1310]" italics="true" pageId="15" pageNumber="108">D. bathyvela</emphasis>
<bibRefCitation id="EF90DB890246FF8D229CFAF8CAB2144C" author="Martynov, A. &amp; Fujiwara, Y. &amp; Tsuchida, S. &amp; Nakano, R. &amp; Sanamyan, N. &amp; Sanamyan, K. &amp; Fletcher, K. &amp; Korshunova, T." journalOrPublisher="Zootaxa" pageId="15" pageNumber="108" pagination="495 - 513" part="4747" refId="ref30619" refString="Martynov, A., Fujiwara, Y., Tsuchida, S., Nakano, R., Sanamyan, N., Sanamyan, K., Fletcher, K. &amp; Korshunova, T. (2020 a) Three new species of the genus Dendronotus from Japan and Russia (Mollusca, Nudibranchia). Zootaxa, 4747, 495 - 513." title="Three new species of the genus Dendronotus from Japan and Russia (Mollusca, Nudibranchia)" type="journal article" year="2020">Martynov et al., 2020a</bibRefCitation>
</taxonomicName>
less specific prey spectrum (including polychaetes and hydroids) has been recorded (
<bibRefCitation id="EF90DB890246FF8D235DFA8DCB311485" author="Roginskaya, I. S." box="[105,295,1392,1418]" editor="Kuznetsov A. P." journalOrPublisher="P. P. Shirshov Institute of Oceanology, Academy of Sciences of the U. S. S. R., Moscow" pageId="15" pageNumber="108" pagination="93 - 110" part="158" refId="ref32127" refString="Roginskaya, I. S. (1990) Additional data on the food and feeding of Dendronotus robustus Verrill, 1879 (Nudibranchia: Dendronotidae) from Dvinsky Bay of the White Sea. In: Kuznetsov A. P. (Eds) Feeding and Bioenergetics of Marine Bottom Invertebrates, pp. 93 - 110, 158. P. P. Shirshov Institute of Oceanology, Academy of Sciences of the U. S. S. R., Moscow." title="Additional data on the food and feeding of Dendronotus robustus Verrill, 1879 (Nudibranchia: Dendronotidae) from Dvinsky Bay of the White Sea" type="book chapter" volumeTitle="Feeding and Bioenergetics of Marine Bottom Invertebrates" year="1990">Roginskaya, 1990</bibRefCitation>
,
<bibRefCitation id="EF90DB890246FF8D2205FA8DCB731485" author="Roginskaya, I. S." box="[305,357,1392,1418]" journalOrPublisher="Opisthobranch Newsl." pageId="15" pageNumber="108" pagination="21 - 23" part="23" refId="ref32213" refString="Roginskaya, I. S. (1997) Predation by a nudibranch Dendronotus robustus from the Sea of Japan on Oweniid polychaetes. Opisthobranch Newsl., 23, 21 - 23." title="Predation by a nudibranch Dendronotus robustus from the Sea of Japan on Oweniid polychaetes" type="journal article" year="1997">1997</bibRefCitation>
).
</paragraph>
<paragraph id="8BBEA6780246FF8D2356FA45CB07176D" blockId="15.[98,610,1464,1634]" pageId="15" pageNumber="108">
<emphasis id="B9757A6A0246FF8D2356FA45CAD714DD" box="[98,193,1464,1490]" italics="true" pageId="15" pageNumber="108">Remarks</emphasis>
. The genus
<taxonomicName id="4C01DDFB0246FF8D227DFA45CBC114DD" authorityName="Alder &amp; Hancock" authorityYear="1845" box="[329,471,1464,1490]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="15" pageNumber="108" phylum="Mollusca" rank="genus">
<emphasis id="B9757A6A0246FF8D227DFA45CBC114DD" box="[329,471,1464,1490]" italics="true" pageId="15" pageNumber="108">Dendronotus</emphasis>
</taxonomicName>
is restricted here since some species have been moved to other genera. See discussions and remarks under the genera
<taxonomicName id="4C01DDFB0246FF8D2275F9D9CBB91731" authority="Korshunova &amp; Bakken &amp; GrØtan &amp; Johnson &amp; Lundin &amp; Martynov, 2021" authorityName="Korshunova &amp; Bakken &amp; GrØtan &amp; Johnson &amp; Lundin &amp; Martynov" authorityYear="2021" box="[321,431,1572,1598]" class="Gastropoda" family="Dendronotidae" genus="Cabangus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Nudibranchia" pageId="15" pageNumber="108" phylum="Mollusca" rank="genus" status="gen. nov.">
<emphasis id="B9757A6A0246FF8D2275F9D9CBB91731" box="[321,431,1572,1598]" italics="true" pageId="15" pageNumber="108">Cabangus</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A246C7110246FF8D228AF9D9C83F1731" box="[446,553,1572,1598]" pageId="15" pageNumber="108" rank="genus">gen. nov.</taxonomicNameLabel>
and
<taxonomicName id="4C01DDFB0246FF8D2356F9B5CB1D176D" authorityName="Baba" authorityYear="1932" box="[98,267,1608,1634]" class="Gastropoda" family="Dendronotidae" genus="Pseudobornella" higherTaxonomySource="CoL" kingdom="Animalia" order="Nudibranchia" pageId="15" pageNumber="108" phylum="Mollusca" rank="genus">
<emphasis id="B9757A6A0246FF8D2356F9B5CB1D176D" box="[98,267,1608,1634]" italics="true" pageId="15" pageNumber="108">Pseudobornella</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="8BBEA6780246FF922356F96DC9221001" blockId="15.[98,609,1679,1742]" lastBlockId="16.[676,1187,171,1742]" lastPageId="16" lastPageNumber="109" pageId="15" pageNumber="108">
<emphasis id="B9757A6A0246FF8D2356F96DCB2017A6" box="[98,310,1680,1706]" italics="true" pageId="15" pageNumber="108">Species composition</emphasis>
. Diagnoses and remarksfor 27 included species are listed below. For each species, an extended diagnosis is provided when possible. The fine-scale diagnostics of the genus
<taxonomicName id="4C01DDFB0246FF8D21C3FF0EC9941002" authorityName="Alder &amp; Hancock" authorityYear="1845" box="[759,898,243,269]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="15" pageNumber="108" phylum="Mollusca" rank="genus">
<emphasis id="B9757A6A0246FF8D21C3FF0EC9941002" box="[759,898,243,269]" italics="true" pageId="15" pageNumber="108">Dendronotus</emphasis>
</taxonomicName>
is a complex task since variation patterns of habitus and anatomical characters overlap in various species. By a careful listing it is possible to step-by-step fulfill the task and present a mutilelevel organismal diversity, though not in a strictly defined form, as was targeted previously (e.g.,
<bibRefCitation id="EF90DB890246FF8D21F2FE13C9481307" author="Odhner, N." box="[710,862,494,520]" journalOrPublisher="Mem. Mus. R. Hist. Nat. Belg." pageId="15" pageNumber="108" pagination="1057 - 1128" part="3" refId="ref31525" refString="Odhner, N. (1936) NudibranchiaDendronotacea - A revision of the system. Mem. Mus. R. Hist. Nat. Belg. (Ser. 2), 3, 1057 - 1128." title="NudibranchiaDendronotacea - A revision of the system" type="journal article" year="1936">Odhner, 1936</bibRefCitation>
;
<bibRefCitation id="EF90DB890246FF8D205BFE13CE341306" author="Robilliard, G." box="[879,1058,494,521]" journalOrPublisher="Veliger" pageId="15" pageNumber="108" pagination="433 - 479" part="12" refId="ref31995" refString="Robilliard, G. (1970) The systematics and some aspects of the ecology of the genus Dendronotus. Veliger, 12, 433 - 479." title="The systematics and some aspects of the ecology of the genus Dendronotus" type="journal article" year="1970">Robilliard, 1970</bibRefCitation>
), but in a complex form that is best suited to the evolutionary mosaic in character evolution. There are also still uncertanties regarding some particular characters as well as previous overinterpretations. First of all, previously the pattern of branching in the dorsolateral appendages was claimed to be of considerable taxonomic importance (
<bibRefCitation id="EF90DB890246FF8D208CFCF3C8A11243" author="Ekimova, I. &amp; Korshunova, T. &amp; Schepetov, D. &amp; Neretina, T. &amp; Sanamyan, N. &amp; Martynov, A." journalOrPublisher="Zool. J. Linn. Soc." pageId="15" pageNumber="108" pagination="841 - 886" part="173" refId="ref28761" refString="Ekimova, I., Korshunova, T., Schepetov, D., Neretina, T., Sanamyan, N. &amp; Martynov, A. (2015) Integrative systematics of northern and Arctic nudibranchs of the genus Dendronotus (Mollusca, Gastropoda), with descriptions of three new species. Zool. J. Linn. Soc., 173, 841 - 886." title="Integrative systematics of northern and Arctic nudibranchs of the genus Dendronotus (Mollusca, Gastropoda), with descriptions of three new species" type="journal article" year="2015">Ekimova et al., 2015</bibRefCitation>
:
<figureCitation id="133ABAFD0246FF8D21F7FCCCC9141243" box="[707,770,817,844]" captionStart="FIGURE 8" captionStartId="24.[129,196,1404,1425]" captionTargetBox="[135,1181,170,1372]" captionTargetId="figure-1@24.[135,1181,170,1372]" captionTargetPageId="24" captionText="FIGURE 8 Dendronotus elegans. External and internal morphology.a: living specimen ZMMU Op-713 from Greenland, near East Atlantic North American coast; b: jaw, same specimen; c; posterior part of radula, general overview,same specimen; d: close up of smooth central teeth, same; e: close up of lateral teeth, same; f:living specimen ZMMU Op-714 (as “D.niveus”) from the White Sea (subarctic); g: jaw, same specimen; h:masticatory edge of the jaws,same; i: posterior part of radula, general overview,same specimen; j: close up of smooth central teeth, same; k: living specimen ZMMU Op-715 from the Laptev Sea (Arctic); l: jaw, same specimen; m: posterior part of radula, close up of smooth central teeth, same; n: lateral teeth, same.o: discoid prostate with numerous alveols (which considerably exceed the considerably underestimated number presented in the first description of D.niveus), same specimen; Scale bars: b, l, 1mm; c: 300 µm; d, hj, m: 100 µm; g: 1000 µm; e, n: 30 µm. PHOTOGRAPHS BY O. ZIMINA (A, K) AND T. KORSHUNOVA (F). SEM MICROGRAPHS BY A. MARTYNOV" figureDoi="http://doi.org/10.5281/zenodo.4623932" httpUri="https://zenodo.org/record/4623932/files/figure.png" pageId="15" pageNumber="108">Fig. 8</figureCitation>
). In this respect, it is necessary to state, that while some trends exist in a few species, importance of these characters were considerably overestimated partly due to the use of preserved specimens in the analysis. This can be illustrated by the
<taxonomicName id="4C01DDFB0246FF8D20EAFC1BCE9212F0" box="[990,1156,997,1023]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="15" pageNumber="108" phylum="Mollusca" rank="species" species="kamchaticus">
<emphasis id="B9757A6A0246FF8D20EAFC1BCE9212F0" box="[990,1156,997,1023]" italics="true" pageId="15" pageNumber="108">D. kamchaticus</emphasis>
</taxonomicName>
case, when according to
<bibRefCitation id="EF90DB890246FF8D20A9FBF4CE94152B" author="Ekimova, I. &amp; Korshunova, T. &amp; Schepetov, D. &amp; Neretina, T. &amp; Sanamyan, N. &amp; Martynov, A." box="[925,1154,1033,1060]" journalOrPublisher="Zool. J. Linn. Soc." pageId="15" pageNumber="108" pagination="841 - 886" part="173" refId="ref28761" refString="Ekimova, I., Korshunova, T., Schepetov, D., Neretina, T., Sanamyan, N. &amp; Martynov, A. (2015) Integrative systematics of northern and Arctic nudibranchs of the genus Dendronotus (Mollusca, Gastropoda), with descriptions of three new species. Zool. J. Linn. Soc., 173, 841 - 886." title="Integrative systematics of northern and Arctic nudibranchs of the genus Dendronotus (Mollusca, Gastropoda), with descriptions of three new species" type="journal article" year="2015">Ekimova et al. (2015</bibRefCitation>
:
<figureCitation id="133ABAFD0246FF8D21B1FBD0C8CB1548" box="[645,733,1069,1095]" captionStart="FIGURE 8" captionStartId="24.[129,196,1404,1425]" captionTargetBox="[135,1181,170,1372]" captionTargetId="figure-1@24.[135,1181,170,1372]" captionTargetPageId="24" captionText="FIGURE 8 Dendronotus elegans. External and internal morphology.a: living specimen ZMMU Op-713 from Greenland, near East Atlantic North American coast; b: jaw, same specimen; c; posterior part of radula, general overview,same specimen; d: close up of smooth central teeth, same; e: close up of lateral teeth, same; f:living specimen ZMMU Op-714 (as “D.niveus”) from the White Sea (subarctic); g: jaw, same specimen; h:masticatory edge of the jaws,same; i: posterior part of radula, general overview,same specimen; j: close up of smooth central teeth, same; k: living specimen ZMMU Op-715 from the Laptev Sea (Arctic); l: jaw, same specimen; m: posterior part of radula, close up of smooth central teeth, same; n: lateral teeth, same.o: discoid prostate with numerous alveols (which considerably exceed the considerably underestimated number presented in the first description of D.niveus), same specimen; Scale bars: b, l, 1mm; c: 300 µm; d, hj, m: 100 µm; g: 1000 µm; e, n: 30 µm. PHOTOGRAPHS BY O. ZIMINA (A, K) AND T. KORSHUNOVA (F). SEM MICROGRAPHS BY A. MARTYNOV" figureDoi="http://doi.org/10.5281/zenodo.4623932" httpUri="https://zenodo.org/record/4623932/files/figure.png" pageId="15" pageNumber="108">Fig. 8D</figureCitation>
),
<taxonomicName id="4C01DDFB0246FF8D21C3FBD3C9B51548" box="[759,931,1069,1095]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="15" pageNumber="108" phylum="Mollusca" rank="species" species="kamchaticus">
<emphasis id="B9757A6A0246FF8D21C3FBD3C9B51548" box="[759,931,1069,1095]" italics="true" pageId="15" pageNumber="108">D. kamchaticus</emphasis>
</taxonomicName>
has bulbous short branches that differ considerably from those in any other
<taxonomicName id="4C01DDFB0246FF8D203AFB88C98F1580" authorityName="Alder &amp; Hancock" authorityYear="1845" box="[782,921,1141,1167]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="15" pageNumber="108" phylum="Mollusca" rank="genus">
<emphasis id="B9757A6A0246FF8D203AFB88C98F1580" box="[782,921,1141,1167]" italics="true" pageId="15" pageNumber="108">Dendronotus</emphasis>
</taxonomicName>
species. This resulted from the use of preserved specimens with considerably contracted branches, whereas the living adult specimens of
<taxonomicName id="4C01DDFB0246FF8D20E9FB1CCE9215F5" box="[989,1156,1248,1274]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="15" pageNumber="108" phylum="Mollusca" rank="species" species="kamchaticus">
<emphasis id="B9757A6A0246FF8D20E9FB1CCE9215F5" box="[989,1156,1248,1274]" italics="true" pageId="15" pageNumber="108">D. kamchaticus</emphasis>
</taxonomicName>
have elongated branches of dorsolateral appendages, which are not fundamentally different from the
<typeStatus id="54BA18DA0246FF8D2068FAB0C99A1468" box="[860,908,1357,1383]" pageId="15" pageNumber="108">type</typeStatus>
species of the genus,
<taxonomicName id="4C01DDFB0246FF8D21B1FA8CCE971484" authority="(Korshunova et al., 2016 a)" baseAuthorityName="Korshunova" baseAuthorityYear="2016" box="[645,1153,1392,1419]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="15" pageNumber="108" phylum="Mollusca" rank="species" species="frondosus">
<emphasis id="B9757A6A0246FF8D21B1FA8CC90C1485" box="[645,794,1392,1418]" italics="true" pageId="15" pageNumber="108">D. frondosus</emphasis>
(
<bibRefCitation id="EF90DB890246FF8D200CFA8DCE6C1484" author="Korshunova, T. &amp; Sanamyan, N. &amp; Zimina, O. &amp; Fletcher, K. &amp; Martynov, A." box="[824,1146,1392,1419]" journalOrPublisher="ZooKeys" pageId="15" pageNumber="108" pagination="19 - 42" part="630" refId="ref30101" refString="Korshunova, T., Sanamyan, N., Zimina, O., Fletcher, K. &amp; Martynov, A. (2016 a) Two new species and a remarkable record of the genus Dendronotus from the North Pacific and Arctic oceans (Nudibranchia). ZooKeys, 630, 19 - 42." title="Two new species and a remarkable record of the genus Dendronotus from the North Pacific and Arctic oceans (Nudibranchia)" type="journal article" year="2016">Korshunova et al., 2016a</bibRefCitation>
)
</taxonomicName>
. The description of
<taxonomicName id="4C01DDFB0246FF8D2059FA68C9CC14A1" authorityName="Martynov, Fujiwara, Tsuchida, R.Nakano, N.Sanamyan, K.Sanamyan, Fletcher &amp; Korshunova" authorityYear="2020" box="[877,986,1428,1454]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="15" pageNumber="108" phylum="Mollusca" rank="species" species="zakuro">
<emphasis id="B9757A6A0246FF8D2059FA68C9CC14A1" box="[877,986,1428,1454]" italics="true" pageId="15" pageNumber="108">D. zakuro</emphasis>
</taxonomicName>
(similar to
<taxonomicName id="4C01DDFB0246FF8D275AFA68C91B14DD" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="15" pageNumber="108" phylum="Mollusca" rank="species" species="kamchaticus">
<emphasis id="B9757A6A0246FF8D275AFA68C91B14DD" italics="true" pageId="15" pageNumber="108">D. kamchaticus</emphasis>
</taxonomicName>
by radula morphology) is based on recently collected specimens with both shorter “bulbous” (see
<bibRefCitation id="EF90DB890246FF8D20B0F9FDCE941715" author="Martynov, A. &amp; Fujiwara, Y. &amp; Tsuchida, S. &amp; Nakano, R. &amp; Sanamyan, N. &amp; Sanamyan, K. &amp; Fletcher, K. &amp; Korshunova, T." box="[900,1154,1536,1562]" journalOrPublisher="Zootaxa" pageId="15" pageNumber="108" pagination="495 - 513" part="4747" refId="ref30619" refString="Martynov, A., Fujiwara, Y., Tsuchida, S., Nakano, R., Sanamyan, N., Sanamyan, K., Fletcher, K. &amp; Korshunova, T. (2020 a) Three new species of the genus Dendronotus from Japan and Russia (Mollusca, Nudibranchia). Zootaxa, 4747, 495 - 513." title="Three new species of the genus Dendronotus from Japan and Russia (Mollusca, Nudibranchia)" type="journal article" year="2020">Martynov et al., 2020a</bibRefCitation>
:
<figureCitation id="133ABAFD0246FF8D21B1F9DEC8C51731" box="[645,723,1571,1598]" captionStart="FIGURE 3" captionStartId="8.[129,196,1542,1563]" captionTargetBox="[129,1187,170,1506]" captionTargetId="figure-1@8.[129,1187,170,1510]" captionTargetPageId="8" captionText="FIGURE 3 A.Map of the Northern Hemisphere showing the distribution of D.elegans (including “D.niveus”), D.kalikal, and D.yrjargul sp. nov.The type locality of D.elegans is indicated by crossed lines. B. The haplotype networks based on cytochrome c oxidase subunit I (COI), 16S rRNA (16S), and 28S rRNA (28S) molecular data showing genetic mutations occurring within D.elegans (“D.niveus”), D. kalikal, and D.yrjargul sp. nov. specimens.Abbreviations:BeSt Bering Strait; BS Barents Sea; Gr Greenland; Km Kamchatka; KS Kara Sea; Ku Kuril Islands; LS Laptev Sea; Nf Newfoundland; NS Norwegian Sea; WS White Sea.Type locality of D.elegans is marked on the map by “x”." figureDoi="http://doi.org/10.5281/zenodo.4623922" httpUri="https://zenodo.org/record/4623922/files/figure.png" pageId="15" pageNumber="108">Fig. 3A</figureCitation>
) and elongate dorsolateral appendages (
<bibRefCitation id="EF90DB890246FF8D21B9F9B5C996176D" author="Martynov, A. &amp; Fujiwara, Y. &amp; Tsuchida, S. &amp; Nakano, R. &amp; Sanamyan, N. &amp; Sanamyan, K. &amp; Fletcher, K. &amp; Korshunova, T." box="[653,896,1607,1634]" journalOrPublisher="Zootaxa" pageId="15" pageNumber="108" pagination="495 - 513" part="4747" refId="ref30619" refString="Martynov, A., Fujiwara, Y., Tsuchida, S., Nakano, R., Sanamyan, N., Sanamyan, K., Fletcher, K. &amp; Korshunova, T. (2020 a) Three new species of the genus Dendronotus from Japan and Russia (Mollusca, Nudibranchia). Zootaxa, 4747, 495 - 513." title="Three new species of the genus Dendronotus from Japan and Russia (Mollusca, Nudibranchia)" type="journal article" year="2020">Martynov et al., 2020a</bibRefCitation>
:
<figureCitation id="133ABAFD0246FF8D20B3F9BAC9FA176D" box="[903,1004,1607,1634]" captionStart="FIGURE 3" captionStartId="8.[129,196,1542,1563]" captionTargetBox="[129,1187,170,1506]" captionTargetId="figure-1@8.[129,1187,170,1510]" captionTargetPageId="8" captionText="FIGURE 3 A.Map of the Northern Hemisphere showing the distribution of D.elegans (including “D.niveus”), D.kalikal, and D.yrjargul sp. nov.The type locality of D.elegans is indicated by crossed lines. B. The haplotype networks based on cytochrome c oxidase subunit I (COI), 16S rRNA (16S), and 28S rRNA (28S) molecular data showing genetic mutations occurring within D.elegans (“D.niveus”), D. kalikal, and D.yrjargul sp. nov. specimens.Abbreviations:BeSt Bering Strait; BS Barents Sea; Gr Greenland; Km Kamchatka; KS Kara Sea; Ku Kuril Islands; LS Laptev Sea; Nf Newfoundland; NS Norwegian Sea; WS White Sea.Type locality of D.elegans is marked on the map by “x”." figureDoi="http://doi.org/10.5281/zenodo.4623922" httpUri="https://zenodo.org/record/4623922/files/figure.png" pageId="15" pageNumber="108">Fig.3D, E</figureCitation>
), whichisdue to differences in physiological conditions. The preserved specimen of
<taxonomicName id="4C01DDFB0246FF8D20B4F96DC9F217A6" authorityName="Martynov, Fujiwara, Tsuchida, R.Nakano, N.Sanamyan, K.Sanamyan, Fletcher &amp; Korshunova" authorityYear="2020" box="[896,996,1679,1705]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="15" pageNumber="108" phylum="Mollusca" rank="species" species="zakuro">
<emphasis id="B9757A6A0246FF8D20B4F96DC9F217A6" box="[896,996,1679,1705]" italics="true" pageId="15" pageNumber="108">D.zakuro</emphasis>
</taxonomicName>
(see
<bibRefCitation id="EF90DB890246FF8D272DF96DC91F17C2" author="Martynov, A. &amp; Fujiwara, Y. &amp; Tsuchida, S. &amp; Nakano, R. &amp; Sanamyan, N. &amp; Sanamyan, K. &amp; Fletcher, K. &amp; Korshunova, T." journalOrPublisher="Zootaxa" pageId="15" pageNumber="108" pagination="495 - 513" part="4747" refId="ref30619" refString="Martynov, A., Fujiwara, Y., Tsuchida, S., Nakano, R., Sanamyan, N., Sanamyan, K., Fletcher, K. &amp; Korshunova, T. (2020 a) Three new species of the genus Dendronotus from Japan and Russia (Mollusca, Nudibranchia). Zootaxa, 4747, 495 - 513." title="Three new species of the genus Dendronotus from Japan and Russia (Mollusca, Nudibranchia)" type="journal article" year="2020">Martynov et al., 2020a</bibRefCitation>
:
<figureCitation id="133ABAFD0246FF8D2025F94EC94B17C2" box="[785,861,1715,1741]" captionStart="FIGURE 3" captionStartId="8.[129,196,1542,1563]" captionTargetBox="[129,1187,170,1506]" captionTargetId="figure-1@8.[129,1187,170,1510]" captionTargetPageId="8" captionText="FIGURE 3 A.Map of the Northern Hemisphere showing the distribution of D.elegans (including “D.niveus”), D.kalikal, and D.yrjargul sp. nov.The type locality of D.elegans is indicated by crossed lines. B. The haplotype networks based on cytochrome c oxidase subunit I (COI), 16S rRNA (16S), and 28S rRNA (28S) molecular data showing genetic mutations occurring within D.elegans (“D.niveus”), D. kalikal, and D.yrjargul sp. nov. specimens.Abbreviations:BeSt Bering Strait; BS Barents Sea; Gr Greenland; Km Kamchatka; KS Kara Sea; Ku Kuril Islands; LS Laptev Sea; Nf Newfoundland; NS Norwegian Sea; WS White Sea.Type locality of D.elegans is marked on the map by “x”." figureDoi="http://doi.org/10.5281/zenodo.4623922" httpUri="https://zenodo.org/record/4623922/files/figure.png" pageId="15" pageNumber="108">Fig. 3C</figureCitation>
) demonstrates very similar “bulbous” branches, which were incorrectly depicted as specific for
<taxonomicName id="4C01DDFB0259FF9222A7FF2DC82C11E6" box="[403,570,207,233]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="109" phylum="Mollusca" rank="species" species="kamchaticus">
<emphasis id="B9757A6A0259FF9222A7FF2DC82C11E6" box="[403,570,207,233]" italics="true" pageId="16" pageNumber="109">D. kamchaticus</emphasis>
</taxonomicName>
in its original description (
<bibRefCitation id="EF90DB890259FF922249FF0EC86B1001" author="Ekimova, I. &amp; Korshunova, T. &amp; Schepetov, D. &amp; Neretina, T. &amp; Sanamyan, N. &amp; Martynov, A." box="[381,637,243,270]" journalOrPublisher="Zool. J. Linn. Soc." pageId="16" pageNumber="109" pagination="841 - 886" part="173" refId="ref28761" refString="Ekimova, I., Korshunova, T., Schepetov, D., Neretina, T., Sanamyan, N. &amp; Martynov, A. (2015) Integrative systematics of northern and Arctic nudibranchs of the genus Dendronotus (Mollusca, Gastropoda), with descriptions of three new species. Zool. J. Linn. Soc., 173, 841 - 886." title="Integrative systematics of northern and Arctic nudibranchs of the genus Dendronotus (Mollusca, Gastropoda), with descriptions of three new species" type="journal article" year="2015">Ekimova et al., 2015</bibRefCitation>
:
<figureCitation id="133ABAFD0259FF9223B5FEEACACE103E" box="[129,216,279,305]" captionStart="FIGURE 8" captionStartId="24.[129,196,1404,1425]" captionTargetBox="[135,1181,170,1372]" captionTargetId="figure-1@24.[135,1181,170,1372]" captionTargetPageId="24" captionText="FIGURE 8 Dendronotus elegans. External and internal morphology.a: living specimen ZMMU Op-713 from Greenland, near East Atlantic North American coast; b: jaw, same specimen; c; posterior part of radula, general overview,same specimen; d: close up of smooth central teeth, same; e: close up of lateral teeth, same; f:living specimen ZMMU Op-714 (as “D.niveus”) from the White Sea (subarctic); g: jaw, same specimen; h:masticatory edge of the jaws,same; i: posterior part of radula, general overview,same specimen; j: close up of smooth central teeth, same; k: living specimen ZMMU Op-715 from the Laptev Sea (Arctic); l: jaw, same specimen; m: posterior part of radula, close up of smooth central teeth, same; n: lateral teeth, same.o: discoid prostate with numerous alveols (which considerably exceed the considerably underestimated number presented in the first description of D.niveus), same specimen; Scale bars: b, l, 1mm; c: 300 µm; d, hj, m: 100 µm; g: 1000 µm; e, n: 30 µm. PHOTOGRAPHS BY O. ZIMINA (A, K) AND T. KORSHUNOVA (F). SEM MICROGRAPHS BY A. MARTYNOV" figureDoi="http://doi.org/10.5281/zenodo.4623932" httpUri="https://zenodo.org/record/4623932/files/figure.png" pageId="16" pageNumber="109">Fig. 8D</figureCitation>
) (see details in Synopsis under
<taxonomicName id="4C01DDFB0259FF92215DFEE5CB1F105A" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="109" phylum="Mollusca" rank="species" species="kamchaticus">
<emphasis id="B9757A6A0259FF92215DFEE5CB1F105A" italics="true" pageId="16" pageNumber="109">D. kamchaticus</emphasis>
</taxonomicName>
).
<taxonomicName id="4C01DDFB0259FF922215FEC6CBE9105A" box="[289,511,315,341]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="109" phylum="Mollusca" rank="species" species="lacteus">
<emphasis id="B9757A6A0259FF922215FEC6CBE9105A" box="[289,511,315,341]" italics="true" pageId="16" pageNumber="109">Dendronotus lacteus</emphasis>
</taxonomicName>
apparently has shorter tertiary branches, and a “curly” appearance of the dorsolateral appendages. However, variation in
<taxonomicName id="4C01DDFB0259FF922243FE55CBC910CE" box="[375,479,423,449]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="109" phylum="Mollusca" rank="species" species="lacteus">
<emphasis id="B9757A6A0259FF922243FE55CBC910CE" box="[375,479,423,449]" italics="true" pageId="16" pageNumber="109">D. lacteus</emphasis>
</taxonomicName>
does not only concern ontogenetic stages, but also occurs among different adult specimens. So, some large specimens of
<taxonomicName id="4C01DDFB0259FF922269FDEECBD11323" box="[349,455,530,556]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="109" phylum="Mollusca" rank="species" species="lacteus">
<emphasis id="B9757A6A0259FF922269FDEECBD11323" box="[349,455,530,556]" italics="true" pageId="16" pageNumber="109">D. lacteus</emphasis>
</taxonomicName>
have quite long elongate branches of dorsolateral appendages, similar to those in
<taxonomicName id="4C01DDFB0259FF92226BFDA6CB361397" authority="(Korshunova et al., 2017 a)" baseAuthorityName="Korshunova" baseAuthorityYear="2017" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="109" phylum="Mollusca" rank="species" species="frondosus">
<emphasis id="B9757A6A0259FF92226BFDA6CBFE137B" box="[351,488,602,628]" italics="true" pageId="16" pageNumber="109">D. frondosus</emphasis>
(
<bibRefCitation id="EF90DB890259FF9222CDFDA7CB0F1397" author="Korshunova, T. &amp; Martynov, A. &amp; Bakken, T. &amp; Evertsen, J. &amp; Fletcher, K. &amp; Mudianta, I. &amp; Saito, H. &amp; Lundin, K. &amp; Schrodl, M. &amp; Picton, B." journalOrPublisher="ZooKeys" pageId="16" pageNumber="109" pagination="1 - 139" part="717" refId="ref29816" refString="Korshunova, T., Martynov, A., Bakken, T., Evertsen, J., Fletcher, K., Mudianta, I., Saito, H., Lundin, K., Schrodl, M. &amp; Picton, B. (2017 a) Polyphyly of the traditional family Flabellinidae affects a major group of Nudibranchia: aeolidacean taxonomic reassessment with descriptions of several new families, genera, and species (Mollusca, Gastropoda). ZooKeys, 717, 1 - 139." title="Polyphyly of the traditional family Flabellinidae affects a major group of Nudibranchia: aeolidacean taxonomic reassessment with descriptions of several new families, genera, and species (Mollusca, Gastropoda)" type="journal article" year="2017">Korshunova et al., 2017a</bibRefCitation>
)
</taxonomicName>
. Most species of the genus
<taxonomicName id="4C01DDFB0259FF9223B5FD5FCB1A13B3" authorityName="Alder &amp; Hancock" authorityYear="1845" box="[129,268,674,700]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="109" phylum="Mollusca" rank="genus">
<emphasis id="B9757A6A0259FF9223B5FD5FCB1A13B3" box="[129,268,674,700]" italics="true" pageId="16" pageNumber="109">Dendronotus</emphasis>
</taxonomicName>
possess considerably branched dorsolateralappendages with distinct tertiary branches (
<bibRefCitation id="EF90DB890259FF922238FD17CBFD120B" author="MacFarland, F. M." box="[268,491,746,772]" journalOrPublisher="Mem. Calif. Acad. Sci." pageId="16" pageNumber="109" pagination="1 - 546" part="6" refId="ref30430" refString="MacFarland, F. M. (1966) Studies of opisthobranchiate mollusks of the Pacific coast of North America. Mem. Calif. Acad. Sci., 6, 1 - 546." title="Studies of opisthobranchiate mollusks of the Pacific coast of North America" type="journal article" year="1966">MacFarland, 1966</bibRefCitation>
;
<bibRefCitation id="EF90DB890259FF92213AFD17CAA41227" author="Robilliard, G." journalOrPublisher="Veliger" pageId="16" pageNumber="109" pagination="433 - 479" part="12" refId="ref31995" refString="Robilliard, G. (1970) The systematics and some aspects of the ecology of the genus Dendronotus. Veliger, 12, 433 - 479." title="The systematics and some aspects of the ecology of the genus Dendronotus" type="journal article" year="1970">Robilliard, 1970</bibRefCitation>
;
<bibRefCitation id="EF90DB890259FF9223FEFCF3CBE91227" author="Korshunova, T. &amp; Sanamyan, N. &amp; Zimina, O. &amp; Fletcher, K. &amp; Martynov, A." box="[202,511,782,808]" journalOrPublisher="ZooKeys" pageId="16" pageNumber="109" pagination="19 - 42" part="630" refId="ref30101" refString="Korshunova, T., Sanamyan, N., Zimina, O., Fletcher, K. &amp; Martynov, A. (2016 a) Two new species and a remarkable record of the genus Dendronotus from the North Pacific and Arctic oceans (Nudibranchia). ZooKeys, 630, 19 - 42." title="Two new species and a remarkable record of the genus Dendronotus from the North Pacific and Arctic oceans (Nudibranchia)" type="journal article" year="2016">Korshunovaet al., 2016a</bibRefCitation>
, b,
<bibRefCitation id="EF90DB890259FF92210BFCF3C86B1227" author="Korshunova, T. &amp; Martynov, A. &amp; Bakken, T. &amp; Evertsen, J. &amp; Fletcher, K. &amp; Mudianta, I. &amp; Saito, H. &amp; Lundin, K. &amp; Schrodl, M. &amp; Picton, B." box="[575,637,782,808]" journalOrPublisher="ZooKeys" pageId="16" pageNumber="109" pagination="1 - 139" part="717" refId="ref29816" refString="Korshunova, T., Martynov, A., Bakken, T., Evertsen, J., Fletcher, K., Mudianta, I., Saito, H., Lundin, K., Schrodl, M. &amp; Picton, B. (2017 a) Polyphyly of the traditional family Flabellinidae affects a major group of Nudibranchia: aeolidacean taxonomic reassessment with descriptions of several new families, genera, and species (Mollusca, Gastropoda). ZooKeys, 717, 1 - 139." title="Polyphyly of the traditional family Flabellinidae affects a major group of Nudibranchia: aeolidacean taxonomic reassessment with descriptions of several new families, genera, and species (Mollusca, Gastropoda)" type="journal article" year="2017">2017a</bibRefCitation>
,
<bibRefCitation id="EF90DB890259FF9223B5FCCFCAD61243" author="Korshunova, T. &amp; Nakano, R. &amp; Fletcher, K. &amp; Sanamyan, N. &amp; Martynov, A." box="[129,192,818,844]" journalOrPublisher="Venus" pageId="16" pageNumber="109" pagination="1 - 14" part="77" refId="ref29910" refString="Korshunova, T., Nakano, R., Fletcher, K., Sanamyan, N. &amp; Martynov, A. (2019 b) First molecular confirmation of the presence of Dendronotus primorjensis Martynov, Sanamyan &amp; Korshunova, 2015 in Japan and new distributional records of Dendronotus species in the North Pacic (Nudibranchia: Dendronotidae). Venus, 77, 1 - 14." title="First molecular confirmation of the presence of Dendronotus primorjensis Martynov, Sanamyan &amp; Korshunova, 2015 in Japan and new distributional records of Dendronotus species in the North Pacic (Nudibranchia: Dendronotidae)" type="journal article" year="2019">2019b</bibRefCitation>
;
<bibRefCitation id="EF90DB890259FF9223E6FCCFCBC81243" author="Martynov, A. &amp; Fujiwara, Y. &amp; Tsuchida, S. &amp; Nakano, R. &amp; Sanamyan, N. &amp; Sanamyan, K. &amp; Fletcher, K. &amp; Korshunova, T." box="[210,478,818,844]" journalOrPublisher="Zootaxa" pageId="16" pageNumber="109" pagination="495 - 513" part="4747" refId="ref30619" refString="Martynov, A., Fujiwara, Y., Tsuchida, S., Nakano, R., Sanamyan, N., Sanamyan, K., Fletcher, K. &amp; Korshunova, T. (2020 a) Three new species of the genus Dendronotus from Japan and Russia (Mollusca, Nudibranchia). Zootaxa, 4747, 495 - 513." title="Three new species of the genus Dendronotus from Japan and Russia (Mollusca, Nudibranchia)" type="journal article" year="2020">Martynov et al., 2020a</bibRefCitation>
). Unbranched dorsolateralappendageswererecentlyreported only for
<taxonomicName id="4C01DDFB0259FF9223EFFC87CB55129C" box="[219,323,889,915]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="109" phylum="Mollusca" rank="species" species="claugei">
<emphasis id="B9757A6A0259FF9223EFFC87CB55129C" box="[219,323,889,915]" italics="true" pageId="16" pageNumber="109">D. claugei</emphasis>
</taxonomicName>
from the deep-sea (
<bibRefCitation id="EF90DB890259FF92212FFC84CACB12B8" author="Valdes, A. &amp; Lundsten, L. &amp; Wilson, N." journalOrPublisher="Zootaxa" pageId="16" pageNumber="109" pagination="401 - 433" part="4" refId="ref32970" refString="Valdes, A., Lundsten, L. &amp; Wilson, N. (2018) Five new deep-sea species of nudibranchs (Gastropoda: Heterobranchia: Cladobranchia) from the Northeast Pacific. Zootaxa, 4526, 4, 401 - 433." title="Five new deep-sea species of nudibranchs (Gastropoda: Heterobranchia: Cladobranchia) from the Northeast Pacific" type="journal article" year="2018">Valdes et al., 2018</bibRefCitation>
), but due to the most basal position of this species (fig. 1) the generic position of
<taxonomicName id="4C01DDFB0259FF9223B5FC1BCAF912F0" box="[129,239,997,1023]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="109" phylum="Mollusca" rank="species" species="claugei">
<emphasis id="B9757A6A0259FF9223B5FC1BCAF912F0" box="[129,239,997,1023]" italics="true" pageId="16" pageNumber="109">D. claugei</emphasis>
</taxonomicName>
needs to be evaluated with more data. For another deep-sea species,
<taxonomicName id="4C01DDFB0259FF92213AFBF7C86C152C" authorityName="Valdes &amp; Bouchet" authorityYear="1998" box="[526,634,1034,1059]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="109" phylum="Mollusca" rank="species" species="comteti">
<emphasis id="B9757A6A0259FF92213AFBF7C86C152C" box="[526,634,1034,1059]" italics="true" pageId="16" pageNumber="109">D. comteti</emphasis>
</taxonomicName>
, the unbranched dorsolateral appendages were also indicated (
<bibRefCitation id="EF90DB890259FF922244FBACC8631564" author="Valdes, A. &amp; Lundsten, L. &amp; Wilson, N." box="[368,629,1105,1131]" journalOrPublisher="Zootaxa" pageId="16" pageNumber="109" pagination="401 - 433" part="4" refId="ref32970" refString="Valdes, A., Lundsten, L. &amp; Wilson, N. (2018) Five new deep-sea species of nudibranchs (Gastropoda: Heterobranchia: Cladobranchia) from the Northeast Pacific. Zootaxa, 4526, 4, 401 - 433." title="Five new deep-sea species of nudibranchs (Gastropoda: Heterobranchia: Cladobranchia) from the Northeast Pacific" type="journal article" year="2018">Valdes et al., 2018: 413</bibRefCitation>
), however, according to the original description of
<taxonomicName id="4C01DDFB0259FF922395FB64CB1815BD" authorityName="Valdes &amp; Bouchet" authorityYear="1998" box="[161,270,1177,1202]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="109" phylum="Mollusca" rank="species" species="comteti">
<emphasis id="B9757A6A0259FF922395FB64CB1815BD" box="[161,270,1177,1202]" italics="true" pageId="16" pageNumber="109">D. comteti</emphasis>
</taxonomicName>
, it possesses both branched and unbrancheddorsolateralappendages (
<bibRefCitation id="EF90DB890259FF92212BFB40CB4315F4" author="Valdes, A. &amp; Bouchet, P." journalOrPublisher="Deep-sea Res. Pt" pageId="16" pageNumber="109" pagination="319 - 327" part="45" refId="ref32934" refString="Valdes, A. &amp; Bouchet, P. (1998) Naked in toxic fluids: a nudibranch mollusc from hydrothermal vents. Deep-sea Res. Pt II, 45, 319 - 327." title="Naked in toxic fluids: a nudibranch mollusc from hydrothermal vents" type="journal article" year="1998">Valdes &amp; Bouchet, 1998: 320</bibRefCitation>
). Unbranched dorsolateral appendagescanoccurinposteriorpartsofsome other
<taxonomicName id="4C01DDFB0259FF9223F4FAD5CB5D144D" authorityName="Alder &amp; Hancock" authorityYear="1845" box="[192,331,1320,1346]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="109" phylum="Mollusca" rank="genus">
<emphasis id="B9757A6A0259FF9223F4FAD5CB5D144D" box="[192,331,1320,1346]" italics="true" pageId="16" pageNumber="109">Dendronotus</emphasis>
</taxonomicName>
species (e.g., in
<taxonomicName id="4C01DDFB0259FF9222C2FAD4C86C144D" authorityName="Korshunova, Sanamyan, Zimina, Fletcher, Martynov" authorityYear="2016" box="[502,634,1320,1346]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="109" phylum="Mollusca" rank="species" species="robilliardi">
<emphasis id="B9757A6A0259FF9222C2FAD4C86C144D" box="[502,634,1320,1346]" italics="true" pageId="16" pageNumber="109">D.robilliardi</emphasis>
</taxonomicName>
, see
<bibRefCitation id="EF90DB890259FF92239DFAB1CBA41468" author="Korshunova, T. &amp; Sanamyan, N. &amp; Zimina, O. &amp; Fletcher, K. &amp; Martynov, A." box="[169,434,1356,1383]" journalOrPublisher="ZooKeys" pageId="16" pageNumber="109" pagination="19 - 42" part="630" refId="ref30101" refString="Korshunova, T., Sanamyan, N., Zimina, O., Fletcher, K. &amp; Martynov, A. (2016 a) Two new species and a remarkable record of the genus Dendronotus from the North Pacific and Arctic oceans (Nudibranchia). ZooKeys, 630, 19 - 42." title="Two new species and a remarkable record of the genus Dendronotus from the North Pacific and Arctic oceans (Nudibranchia)" type="journal article" year="2016">Korshunova et al., 2016a</bibRefCitation>
). For some species, e.g., for
<taxonomicName id="4C01DDFB0259FF9223E9FA8CCB3A1485" authorityName="sensu Bergh" authorityYear="1886" box="[221,300,1392,1418]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="109" phylum="Mollusca" rank="species" species="dalli">
<emphasis id="B9757A6A0259FF9223E9FA8CCB3A1485" box="[221,300,1392,1418]" italics="true" pageId="16" pageNumber="109">D. dalli</emphasis>
</taxonomicName>
in
<bibRefCitation id="EF90DB890259FF922262FA8DC86A1484" author="Ekimova, I. &amp; Korshunova, T. &amp; Schepetov, D. &amp; Neretina, T. &amp; Sanamyan, N. &amp; Martynov, A." box="[342,636,1392,1419]" journalOrPublisher="Zool. J. Linn. Soc." pageId="16" pageNumber="109" pagination="841 - 886" part="173" refId="ref28761" refString="Ekimova, I., Korshunova, T., Schepetov, D., Neretina, T., Sanamyan, N. &amp; Martynov, A. (2015) Integrative systematics of northern and Arctic nudibranchs of the genus Dendronotus (Mollusca, Gastropoda), with descriptions of three new species. Zool. J. Linn. Soc., 173, 841 - 886." title="Integrative systematics of northern and Arctic nudibranchs of the genus Dendronotus (Mollusca, Gastropoda), with descriptions of three new species" type="journal article" year="2015">Ekimova et al. (2015: 862)</bibRefCitation>
, the indication of absent tertiary branches of dorsolateral appendages is incorrect.
<taxonomicName id="4C01DDFB0259FF9223B5FA21CB4214F9" authorityName="sensu Bergh" authorityYear="1886" box="[129,340,1500,1526]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="109" phylum="Mollusca" rank="species" species="dalli">
<emphasis id="B9757A6A0259FF9223B5FA21CB4214F9" box="[129,340,1500,1526]" italics="true" pageId="16" pageNumber="109">Dendronotusdalli</emphasis>
</taxonomicName>
definitely has tertiary branches (
<bibRefCitation id="EF90DB890259FF9223C1F9FDCBCE1714" author="Robilliard, G." box="[245,472,1536,1563]" journalOrPublisher="Veliger" pageId="16" pageNumber="109" pagination="433 - 479" part="12" refId="ref31995" refString="Robilliard, G. (1970) The systematics and some aspects of the ecology of the genus Dendronotus. Veliger, 12, 433 - 479." title="The systematics and some aspects of the ecology of the genus Dendronotus" type="journal article" year="1970">Robilliard, 1970: 453</bibRefCitation>
; presentstudy, fig. 7). Instead, in the genus
<taxonomicName id="4C01DDFB0259FF922285F9D9C8401731" authorityName="Baba" authorityYear="1932" box="[433,598,1572,1598]" class="Gastropoda" family="Dendronotidae" genus="Pseudobornella" higherTaxonomySource="CoL" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="109" phylum="Mollusca" rank="genus">
<emphasis id="B9757A6A0259FF922285F9D9C8401731" box="[433,598,1572,1598]" italics="true" pageId="16" pageNumber="109">Pseudobornella</emphasis>
</taxonomicName>
the generalappearanceof dorsolateralappendages does not show as branches, but it is ctenidiumlike, because of very indistinct (or truly absent) tertiary branches (fig. 2). More details about the genus
<taxonomicName id="4C01DDFB0259FF92202CFF56C9AB11CA" authorityName="Baba" authorityYear="1932" box="[792,957,171,197]" class="Gastropoda" family="Dendronotidae" genus="Pseudobornella" higherTaxonomySource="CoL" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="109" phylum="Mollusca" rank="genus">
<emphasis id="B9757A6A0259FF92202CFF56C9AB11CA" box="[792,957,171,197]" italics="true" pageId="16" pageNumber="109">Pseudobornella</emphasis>
</taxonomicName>
are mentioned after the diagnoses of the species of the genus
<taxonomicName id="4C01DDFB0259FF922190FF0EC9391002" authorityName="Alder &amp; Hancock" authorityYear="1845" box="[676,815,243,269]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="109" phylum="Mollusca" rank="genus">
<emphasis id="B9757A6A0259FF922190FF0EC9391002" box="[676,815,243,269]" italics="true" pageId="16" pageNumber="109">Dendronotus</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="8BBEA6780259FF9221F6FEEAC9791715" blockId="16.[676,1187,171,1742]" pageId="16" pageNumber="109">
There are also various uncertainties and misinterpretations regarding internal characters that need to be addressed before diagnoses can be provided. For example,
<bibRefCitation id="EF90DB890259FF922700FE7EC8F110CE" author="Robilliard, G." journalOrPublisher="Veliger" pageId="16" pageNumber="109" pagination="433 - 479" part="12" refId="ref31995" refString="Robilliard, G. (1970) The systematics and some aspects of the ecology of the genus Dendronotus. Veliger, 12, 433 - 479." title="The systematics and some aspects of the ecology of the genus Dendronotus" type="journal article" year="1970">Robilliard (1970)</bibRefCitation>
suggested to apply several coefficients of jaw proportions. However, it is easy to incorrectlyapply respective angleof measurements and receive very different results, as for example in
<bibRefCitation id="EF90DB890259FF922004FDCBCE30135E" author="Ekimova, I. &amp; Korshunova, T. &amp; Schepetov, D. &amp; Neretina, T. &amp; Sanamyan, N. &amp; Martynov, A." box="[816,1062,566,593]" journalOrPublisher="Zool. J. Linn. Soc." pageId="16" pageNumber="109" pagination="841 - 886" part="173" refId="ref28761" refString="Ekimova, I., Korshunova, T., Schepetov, D., Neretina, T., Sanamyan, N. &amp; Martynov, A. (2015) Integrative systematics of northern and Arctic nudibranchs of the genus Dendronotus (Mollusca, Gastropoda), with descriptions of three new species. Zool. J. Linn. Soc., 173, 841 - 886." title="Integrative systematics of northern and Arctic nudibranchs of the genus Dendronotus (Mollusca, Gastropoda), with descriptions of three new species" type="journal article" year="2015">Ekimova et al. (2015)</bibRefCitation>
compared to the same species which jaws by
<bibRefCitation id="EF90DB890259FF922700FDA7C8FC1397" author="Robilliard, G." journalOrPublisher="Veliger" pageId="16" pageNumber="109" pagination="433 - 479" part="12" refId="ref31995" refString="Robilliard, G. (1970) The systematics and some aspects of the ecology of the genus Dendronotus. Veliger, 12, 433 - 479." title="The systematics and some aspects of the ecology of the genus Dendronotus" type="journal article" year="1970">Robilliard (1970)</bibRefCitation>
. As an example, for
<taxonomicName id="4C01DDFB0259FF9220FAFD82CE0A1397" authorityName="sensu Bergh" authorityYear="1886" box="[974,1052,638,664]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="109" phylum="Mollusca" rank="species" species="dalli">
<emphasis id="B9757A6A0259FF9220FAFD82CE0A1397" box="[974,1052,638,664]" italics="true" pageId="16" pageNumber="109">D. dalli</emphasis>
</taxonomicName>
in
<bibRefCitation id="EF90DB890259FF922775FD83C94A13B2" author="Ekimova, I. &amp; Korshunova, T. &amp; Schepetov, D. &amp; Neretina, T. &amp; Sanamyan, N. &amp; Martynov, A." journalOrPublisher="Zool. J. Linn. Soc." pageId="16" pageNumber="109" pagination="841 - 886" part="173" refId="ref28761" refString="Ekimova, I., Korshunova, T., Schepetov, D., Neretina, T., Sanamyan, N. &amp; Martynov, A. (2015) Integrative systematics of northern and Arctic nudibranchs of the genus Dendronotus (Mollusca, Gastropoda), with descriptions of three new species. Zool. J. Linn. Soc., 173, 841 - 886." title="Integrative systematics of northern and Arctic nudibranchs of the genus Dendronotus (Mollusca, Gastropoda), with descriptions of three new species" type="journal article" year="2015">Ekimova et al. (2015: 863)</bibRefCitation>
an angle of dorsal processes to the longitudinal axis of 25° was indicated, whereas in
<bibRefCitation id="EF90DB890259FF922017FD17CE18120A" author="Robilliard, G." box="[803,1038,746,773]" journalOrPublisher="Veliger" pageId="16" pageNumber="109" pagination="433 - 479" part="12" refId="ref31995" refString="Robilliard, G. (1970) The systematics and some aspects of the ecology of the genus Dendronotus. Veliger, 12, 433 - 479." title="The systematics and some aspects of the ecology of the genus Dendronotus" type="journal article" year="1970">Robilliard (1970: 454)</bibRefCitation>
this was 60°. In the present study we re-checked jaw proportions andpresented updated values for the majority of species with sufficient available information. This character still needs further testing, but some characteristic patterns for different species are likely to occur. Similarly, the number of the prostatic alveoles once was thought to be a character that well characterizes different species (
<bibRefCitation id="EF90DB890259FF9220DDFBD0CE891548" author="Robilliard, G." box="[1001,1183,1069,1095]" journalOrPublisher="Veliger" pageId="16" pageNumber="109" pagination="433 - 479" part="12" refId="ref31995" refString="Robilliard, G. (1970) The systematics and some aspects of the ecology of the genus Dendronotus. Veliger, 12, 433 - 479." title="The systematics and some aspects of the ecology of the genus Dendronotus" type="journal article" year="1970">Robilliard, 1970</bibRefCitation>
;
<bibRefCitation id="EF90DB890259FF922190FBACC9981563" author="Ekimova, I. &amp; Schepetov, D. &amp; Chichvarkhina, O. &amp; Chichvarkhin, A." box="[676,910,1105,1132]" journalOrPublisher="Invertebr. Zool." pageId="16" pageNumber="109" pagination="15 - 42" part="13" refId="ref28832" refString="Ekimova, I., Schepetov, D., Chichvarkhina, O. &amp; Chichvarkhin, A. (2016) Nudibranch molluscs of the genus Dendronotus Alder et Hancock, 1845 (Heterobranchia: Dendronotina) from Northwestern Sea of Japan with description of a new species. Invertebr. Zool., 13, 15 - 42." title="Nudibranch molluscs of the genus Dendronotus Alder et Hancock, 1845 (Heterobranchia: Dendronotina) from Northwestern Sea of Japan with description of a new species" type="journal article" year="2016">Ekimova et al., 2016</bibRefCitation>
). However, our previous (
<bibRefCitation id="EF90DB890259FF92219FFB88C9A21580" author="Korshunova, T. &amp; Sanamyan, N. &amp; Zimina, O. &amp; Fletcher, K. &amp; Martynov, A." box="[683,948,1141,1167]" journalOrPublisher="ZooKeys" pageId="16" pageNumber="109" pagination="19 - 42" part="630" refId="ref30101" refString="Korshunova, T., Sanamyan, N., Zimina, O., Fletcher, K. &amp; Martynov, A. (2016 a) Two new species and a remarkable record of the genus Dendronotus from the North Pacific and Arctic oceans (Nudibranchia). ZooKeys, 630, 19 - 42." title="Two new species and a remarkable record of the genus Dendronotus from the North Pacific and Arctic oceans (Nudibranchia)" type="journal article" year="2016">Korshunova et al. 2016a</bibRefCitation>
, b) and presentstudy show that this character is much more variable, and while there are clear trends in some species, it should generally be used with care. In the species diagnoses below, differences in shape, length and proportions of the copulative organ (penis) are indicated. While in some species the differences in penial morphology are clear, in other species this can be partly due to different degrees of contraction after preservation. This character needs further investigation.
</paragraph>
<paragraph id="8BBEA6780259FF9321F6F9D9CB0B105A" blockId="16.[676,1187,171,1742]" lastBlockId="17.[98,610,171,1741]" lastPageId="17" lastPageNumber="110" pageId="16" pageNumber="109">
In the presentstudy we show that the shape andsize of the bursa copulatrix, atleastamong several
<taxonomicName id="4C01DDFB0259FF922035F996C998178A" authorityName="Alder &amp; Hancock" authorityYear="1845" box="[769,910,1643,1669]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="109" phylum="Mollusca" rank="genus">
<emphasis id="B9757A6A0259FF922035F996C998178A" box="[769,910,1643,1669]" italics="true" pageId="16" pageNumber="109">Dendronotus</emphasis>
</taxonomicName>
species (e.g., between
<taxonomicName id="4C01DDFB0259FF922190F96DC91917A6" box="[676,783,1679,1705]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="109" phylum="Mollusca" rank="species" species="kalikal">
<emphasis id="B9757A6A0259FF922190F96DC91917A6" box="[676,783,1679,1705]" italics="true" pageId="16" pageNumber="109">D. kalikal</emphasis>
</taxonomicName>
and
<taxonomicName id="4C01DDFB0259FF922062F96DC9C617A6" authorityName="Korshunova &amp; Bakken &amp; GrØtan &amp; Johnson &amp; Lundin &amp; Martynov" authorityYear="2021" box="[854,976,1679,1705]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="109" phylum="Mollusca" rank="species" species="yrjargul">
<emphasis id="B9757A6A0259FF922062F96DC9C617A6" box="[854,976,1679,1705]" italics="true" pageId="16" pageNumber="109">D. yrjargul</emphasis>
</taxonomicName>
), are rather constant and can be used as a reliable diagnostic character. However, the exact position of the commonly very small receptaculum seminis can be very difficult to determine, but despite that, it will be given further attention in the diagnoses below.
</paragraph>
<paragraph id="8BBEA6780258FF9023B5FEA2CB6611E6" blockId="17.[98,610,171,1741]" lastBlockId="18.[129,641,171,1742]" lastPageId="18" lastPageNumber="111" pageId="17" pageNumber="110">
Different terms were applied for the structures on the masticatory edge within different species of the genus
<taxonomicName id="4C01DDFB0258FF9322AFFE5AC83F10CE" authorityName="Alder &amp; Hancock" authorityYear="1845" box="[411,553,423,449]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="17" pageNumber="110" phylum="Mollusca" rank="genus">
<emphasis id="B9757A6A0258FF9322AFFE5AC83F10CE" box="[411,553,423,449]" italics="true" pageId="17" pageNumber="110">Dendronotus</emphasis>
</taxonomicName>
. For example,
<bibRefCitation id="EF90DB890258FF9323E5FE36CBB210E9" author="MacFarland, F. M." box="[209,420,459,486]" journalOrPublisher="Mem. Calif. Acad. Sci." pageId="17" pageNumber="110" pagination="1 - 546" part="6" refId="ref30430" refString="MacFarland, F. M. (1966) Studies of opisthobranchiate mollusks of the Pacific coast of North America. Mem. Calif. Acad. Sci., 6, 1 - 546." title="Studies of opisthobranchiate mollusks of the Pacific coast of North America" type="journal article" year="1966">MacFarland (1966)</bibRefCitation>
used “denticles” or “ridge-like” denticles and
<bibRefCitation id="EF90DB890258FF932293FE12C8771306" author="Robilliard, G." box="[423,609,495,521]" journalOrPublisher="Veliger" pageId="17" pageNumber="110" pagination="433 - 479" part="12" refId="ref31995" refString="Robilliard, G. (1970) The systematics and some aspects of the ecology of the genus Dendronotus. Veliger, 12, 433 - 479." title="The systematics and some aspects of the ecology of the genus Dendronotus" type="journal article" year="1970">Robilliard (1970)</bibRefCitation>
applied “denticles”, whereas
<bibRefCitation id="EF90DB890258FF932281FDEFCAB0135E" author="Ekimova, I. &amp; Korshunova, T. &amp; Schepetov, D. &amp; Neretina, T. &amp; Sanamyan, N. &amp; Martynov, A." journalOrPublisher="Zool. J. Linn. Soc." pageId="17" pageNumber="110" pagination="841 - 886" part="173" refId="ref28761" refString="Ekimova, I., Korshunova, T., Schepetov, D., Neretina, T., Sanamyan, N. &amp; Martynov, A. (2015) Integrative systematics of northern and Arctic nudibranchs of the genus Dendronotus (Mollusca, Gastropoda), with descriptions of three new species. Zool. J. Linn. Soc., 173, 841 - 886." title="Integrative systematics of northern and Arctic nudibranchs of the genus Dendronotus (Mollusca, Gastropoda), with descriptions of three new species" type="journal article" year="2015">Ekimova et al. (2015)</bibRefCitation>
used “rodlets”. In this respect, the term “rodlets” was used for rod-like structures that constitute the body of jaws in other nudibranch groups. In the genus
<taxonomicName id="4C01DDFB0258FF932296FD5FC82613B3" authorityName="Alder &amp; Hancock" authorityYear="1845" box="[418,560,674,700]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="17" pageNumber="110" phylum="Mollusca" rank="genus">
<emphasis id="B9757A6A0258FF932296FD5FC82613B3" box="[418,560,674,700]" italics="true" pageId="17" pageNumber="110">Dendronotus</emphasis>
</taxonomicName>
, the elements on the masticatory edges are usually termed as “denticles” when the bodies of jaws are entire. However, in the genus
<taxonomicName id="4C01DDFB0258FF9322E7FCF3C8771227" authorityName="Alder &amp; Hancock" authorityYear="1845" box="[467,609,782,808]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="17" pageNumber="110" phylum="Mollusca" rank="genus">
<emphasis id="B9757A6A0258FF9322E7FCF3C8771227" box="[467,609,782,808]" italics="true" pageId="17" pageNumber="110">Dendronotus</emphasis>
</taxonomicName>
there is a special set of structures in addition to the denticles that appears as broad semidisks or ridges, which can additionally be covered with smaller spine- or denticle-like structures. It is likely that most species of the genus
<taxonomicName id="4C01DDFB0258FF93239DFC18CB2112F0" authorityName="Alder &amp; Hancock" authorityYear="1845" box="[169,311,997,1023]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="17" pageNumber="110" phylum="Mollusca" rank="genus">
<emphasis id="B9757A6A0258FF93239DFC18CB2112F0" box="[169,311,997,1023]" italics="true" pageId="17" pageNumber="110">Dendronotus</emphasis>
</taxonomicName>
possess such structures. In addition to the semi-disks/ridges at the masticatory edges, more normallylooking denticles can occur. Transitive denticles on masticatory edges werealsooccasionally reported forjuveniles of
<taxonomicName id="4C01DDFB0258FF932389FB67CB5015BC" box="[189,326,1177,1203]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="17" pageNumber="110" phylum="Mollusca" rank="species" sensu="lato" species="frondosus">
<emphasis id="B9757A6A0258FF932389FB67CB5015BC" box="[189,326,1177,1203]" italics="true" pageId="17" pageNumber="110">D. frondosus</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A246C7110258FF932279FB64CB7C15BC" box="[333,362,1177,1203]" pageId="17" pageNumber="110" sensu="lato">s.l.</taxonomicNameLabel>
(
<bibRefCitation id="EF90DB890258FF93224DFB64C81315BC" author="Colgan, N." box="[377,517,1177,1203]" journalOrPublisher="Ir. Nat. J." pageId="17" pageNumber="110" pagination="161 - 204" part="23" refId="ref28519" refString="Colgan, N. (1914) The opisthobranch fauna of the shores and shallow waters of County Dublin. Ir. Nat. J., 23 (8 / 9), 161 - 204." title="The opisthobranch fauna of the shores and shallow waters of County Dublin" type="journal article" year="1914">Colgan, 1914</bibRefCitation>
;
<bibRefCitation id="EF90DB890258FF93213BFB64CB0B15D8" author="Wagele, H. &amp; Willan, R." journalOrPublisher="Zool. J. Linn. Soc." pageId="17" pageNumber="110" pagination="83 - 181" part="130" refId="ref33246" refString="Wagele, H., Willan, R. (2000) Phylogeny of the Nudibranchia. Zool. J. Linn. Soc., 130, 83 - 181." title="Phylogeny of the Nudibranchia" type="journal article" year="2000">Wägele &amp; Willan, 2000</bibRefCitation>
). Therefore, in the present study we use two different terms “ridge-like structures” and “denticles” when characterizing the masticatory edges of
<taxonomicName id="4C01DDFB0258FF9322E7FAD5C877144D" authorityName="Alder &amp; Hancock" authorityYear="1845" box="[467,609,1320,1346]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="17" pageNumber="110" phylum="Mollusca" rank="genus">
<emphasis id="B9757A6A0258FF9322E7FAD5C877144D" box="[467,609,1320,1346]" italics="true" pageId="17" pageNumber="110">Dendronotus</emphasis>
</taxonomicName>
species. The few denticles seen in many cases are likely a precursor that further transits into semi-disk structures on the masicatory edges (see e.g., this clearly for
<taxonomicName id="4C01DDFB0258FF9322BDFA44CBE514DD" box="[393,499,1464,1490]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="17" pageNumber="110" phylum="Mollusca" rank="species" species="kalikal">
<emphasis id="B9757A6A0258FF9322BDFA44CBE514DD" box="[393,499,1464,1490]" italics="true" pageId="17" pageNumber="110">D. kalikal</emphasis>
</taxonomicName>
, fig. 4Q). Because thefull row of denticles and/or ridges is not documented for all species due to difficulties during preparation, the exact number for these structures can be uncertain. Though whenever possible, we already presented the number of denticles/ridges, for example the recently described
<taxonomicName id="4C01DDFB0258FF932208F94ECBDC17C2" authorityName="Alder &amp; Hancock" authorityYear="1845" box="[316,458,1715,1741]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="17" pageNumber="110" phylum="Mollusca" rank="genus">
<emphasis id="B9757A6A0258FF932208F94ECBDC17C2" box="[316,458,1715,1741]" italics="true" pageId="17" pageNumber="110">Dendronotus</emphasis>
</taxonomicName>
species (e.g.,
<bibRefCitation id="EF90DB890258FF9321B1FF51C98911CA" author="Martynov, A. &amp; Fujiwara, Y. &amp; Tsuchida, S. &amp; Nakano, R. &amp; Sanamyan, N. &amp; Sanamyan, K. &amp; Fletcher, K. &amp; Korshunova, T." box="[645,927,171,198]" journalOrPublisher="Zootaxa" pageId="17" pageNumber="110" pagination="495 - 513" part="4747" refId="ref30619" refString="Martynov, A., Fujiwara, Y., Tsuchida, S., Nakano, R., Sanamyan, N., Sanamyan, K., Fletcher, K. &amp; Korshunova, T. (2020 a) Three new species of the genus Dendronotus from Japan and Russia (Mollusca, Nudibranchia). Zootaxa, 4747, 495 - 513." title="Three new species of the genus Dendronotus from Japan and Russia (Mollusca, Nudibranchia)" type="journal article" year="2020">Martynov et al., 2020a</bibRefCitation>
), in the diagnoses below we do not provide the number because it can be misleading, because only a part of the row was counted in various species. True rodlets can occur in addition, as part of the thin labial cuticle, which is separated from the masticatory edges. Such true rodlets have been well documented for the tropical representatives of the family
<taxonomicName id="4C01DDFB0258FF9320ACFE37C9211306" authority="(Pola &amp; Stout, 2008)" baseAuthorityName="Pola &amp; Stout" baseAuthorityYear="2008" class="Gastropoda" family="Dendronotidae" kingdom="Animalia" order="Nudibranchia" pageId="17" pageNumber="110" phylum="Mollusca" rank="family">
Dendronotidae (
<bibRefCitation id="EF90DB890258FF932766FE37C9381306" author="Pola, M. &amp; Stout, C." journalOrPublisher="Zootaxa" pageId="17" pageNumber="110" pagination="45 - 66" part="1960" refId="ref31659" refString="Pola, M. &amp; Stout, C. (2008) Description of the first two tropical Indo-Pacific species of Dendronotus (Gastropoda: Nudibranchia) with new data of the poorly known species Dendronotus gracilis Baba, 1949. Zootaxa, 1960, 45 - 66." title="Description of the first two tropical Indo-Pacific species of Dendronotus (Gastropoda: Nudibranchia) with new data of the poorly known species Dendronotus gracilis Baba, 1949" type="journal article" year="2008">Pola &amp; Stout, 2008</bibRefCitation>
)
</taxonomicName>
, and possibly also occur in many
<taxonomicName id="4C01DDFB0258FF9321E5FDEFC9491323" authorityName="Alder &amp; Hancock" authorityYear="1845" box="[721,863,530,556]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="17" pageNumber="110" phylum="Mollusca" rank="genus">
<emphasis id="B9757A6A0258FF9321E5FDEFC9491323" box="[721,863,530,556]" italics="true" pageId="17" pageNumber="110">Dendronotus</emphasis>
</taxonomicName>
species, since the labial cuticle with rodletswere already described for several representatives (see e.g.,
<bibRefCitation id="EF90DB890258FF9320CDFDA7C8AD1396" author="MacFarland, F. M." journalOrPublisher="Mem. Calif. Acad. Sci." pageId="17" pageNumber="110" pagination="1 - 546" part="6" refId="ref30430" refString="MacFarland, F. M. (1966) Studies of opisthobranchiate mollusks of the Pacific coast of North America. Mem. Calif. Acad. Sci., 6, 1 - 546." title="Studies of opisthobranchiate mollusks of the Pacific coast of North America" type="journal article" year="1966">MacFarland, 1966</bibRefCitation>
). However, these characters remain largely untested by modern methods and we do not include it in the diagnoses below. As an additional complication, denticles and especially ridge-like structures on the masticatory edges can be easily discarded during preparation, and therefore in some previous descriptions it was inappropriately indicated that masticatory edges are devoid of any denticles or ridges. For example, the absence of the denticles on the masticatory edges of
<taxonomicName id="4C01DDFB0258FF93219DFBF7C923152C" box="[681,821,1033,1059]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="17" pageNumber="110" phylum="Mollusca" rank="species" species="frondosus">
<emphasis id="B9757A6A0258FF93219DFBF7C923152C" box="[681,821,1033,1059]" italics="true" pageId="17" pageNumber="110">D. frondosus</emphasis>
</taxonomicName>
were incorrectly mentioned in
<bibRefCitation id="EF90DB890258FF932193FBD0C9B61547" author="Ekimova, I. &amp; Korshunova, T. &amp; Schepetov, D. &amp; Neretina, T. &amp; Sanamyan, N. &amp; Martynov, A." box="[679,928,1069,1096]" journalOrPublisher="Zool. J. Linn. Soc." pageId="17" pageNumber="110" pagination="841 - 886" part="173" refId="ref28761" refString="Ekimova, I., Korshunova, T., Schepetov, D., Neretina, T., Sanamyan, N. &amp; Martynov, A. (2015) Integrative systematics of northern and Arctic nudibranchs of the genus Dendronotus (Mollusca, Gastropoda), with descriptions of three new species. Zool. J. Linn. Soc., 173, 841 - 886." title="Integrative systematics of northern and Arctic nudibranchs of the genus Dendronotus (Mollusca, Gastropoda), with descriptions of three new species" type="journal article" year="2015">Ekimova et al. (2015)</bibRefCitation>
, which was further used as a putative difference between
<taxonomicName id="4C01DDFB0258FF932707FBAFC8D51580" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="17" pageNumber="110" phylum="Mollusca" rank="species" species="frondosus">
<emphasis id="B9757A6A0258FF932707FBAFC8D51580" italics="true" pageId="17" pageNumber="110">D. frondosus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C01DDFB0258FF932038FB8BC8D515BB" authority="(Ekimova et al., 2016)" baseAuthorityName="Ekimova" baseAuthorityYear="2016" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="17" pageNumber="110" phylum="Mollusca" rank="species" species="primorjensis">
<emphasis id="B9757A6A0258FF932038FB8BC9AE1580" box="[780,952,1142,1167]" italics="true" pageId="17" pageNumber="110">D. primorjensis</emphasis>
(
<bibRefCitation id="EF90DB890258FF9320FAFB88C8AA15BB" author="Ekimova, I. &amp; Schepetov, D. &amp; Chichvarkhina, O. &amp; Chichvarkhin, A." journalOrPublisher="Invertebr. Zool." pageId="17" pageNumber="110" pagination="15 - 42" part="13" refId="ref28832" refString="Ekimova, I., Schepetov, D., Chichvarkhina, O. &amp; Chichvarkhin, A. (2016) Nudibranch molluscs of the genus Dendronotus Alder et Hancock, 1845 (Heterobranchia: Dendronotina) from Northwestern Sea of Japan with description of a new species. Invertebr. Zool., 13, 15 - 42." title="Nudibranch molluscs of the genus Dendronotus Alder et Hancock, 1845 (Heterobranchia: Dendronotina) from Northwestern Sea of Japan with description of a new species" type="journal article" year="2016">Ekimova et al., 2016</bibRefCitation>
)
</taxonomicName>
. However, in both species denticles on the masticatory edges are definitely present (
<bibRefCitation id="EF90DB890258FF9321B9FB1DC9BB15F4" author="Korshunova, T. &amp; Sanamyan, N. &amp; Zimina, O. &amp; Fletcher, K. &amp; Martynov, A." box="[653,941,1248,1275]" journalOrPublisher="ZooKeys" pageId="17" pageNumber="110" pagination="19 - 42" part="630" refId="ref30101" refString="Korshunova, T., Sanamyan, N., Zimina, O., Fletcher, K. &amp; Martynov, A. (2016 a) Two new species and a remarkable record of the genus Dendronotus from the North Pacific and Arctic oceans (Nudibranchia). ZooKeys, 630, 19 - 42." title="Two new species and a remarkable record of the genus Dendronotus from the North Pacific and Arctic oceans (Nudibranchia)" type="journal article" year="2016">Korshunova et al., 2016a</bibRefCitation>
). The same is true also for other species, such as
<taxonomicName id="4C01DDFB0258FF9320E8FAF8CE921411" box="[988,1156,1284,1310]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="17" pageNumber="110" phylum="Mollusca" rank="species" species="kamchaticus">
<emphasis id="B9757A6A0258FF9320E8FAF8CE921411" box="[988,1156,1284,1310]" italics="true" pageId="17" pageNumber="110">D. kamchaticus</emphasis>
</taxonomicName>
and
<emphasis id="B9757A6A0258FF932182FAD4C934144D" box="[694,802,1320,1346]" italics="true" pageId="17" pageNumber="110">
<taxonomicName id="4C01DDFB0258FF932182FAD4C908144D" box="[694,798,1320,1346]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="17" pageNumber="110" phylum="Mollusca" rank="species" species="kalikal">D. kalikal</taxonomicName>
.
</emphasis>
For
<taxonomicName id="4C01DDFB0258FF932066FAD4C9EF144D" box="[850,1017,1320,1346]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="17" pageNumber="110" phylum="Mollusca" rank="species" species="kamchaticus">
<emphasis id="B9757A6A0258FF932066FAD4C9EF144D" box="[850,1017,1320,1346]" italics="true" pageId="17" pageNumber="110">D. kamchaticus</emphasis>
</taxonomicName>
the absence of any structures on the mascticatory edges was indicated in the original description. For
<taxonomicName id="4C01DDFB0258FF9321B1FA68C8FC14A1" box="[645,746,1428,1454]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="17" pageNumber="110" phylum="Mollusca" rank="species" species="kalikal">
<emphasis id="B9757A6A0258FF9321B1FA68C8FC14A1" box="[645,746,1428,1454]" italics="true" pageId="17" pageNumber="110">D. kalikal</emphasis>
</taxonomicName>
the presence of merely “rodlets” was reported in the original description (
<bibRefCitation id="EF90DB890258FF932714FA45C91714F8" author="Ekimova, I. &amp; Korshunova, T. &amp; Schepetov, D. &amp; Neretina, T. &amp; Sanamyan, N. &amp; Martynov, A." journalOrPublisher="Zool. J. Linn. Soc." pageId="17" pageNumber="110" pagination="841 - 886" part="173" refId="ref28761" refString="Ekimova, I., Korshunova, T., Schepetov, D., Neretina, T., Sanamyan, N. &amp; Martynov, A. (2015) Integrative systematics of northern and Arctic nudibranchs of the genus Dendronotus (Mollusca, Gastropoda), with descriptions of three new species. Zool. J. Linn. Soc., 173, 841 - 886." title="Integrative systematics of northern and Arctic nudibranchs of the genus Dendronotus (Mollusca, Gastropoda), with descriptions of three new species" type="journal article" year="2015">Ekimova et al., 2015</bibRefCitation>
). However in reality denticles and ridge-like structures are present in both of these species (
<bibRefCitation id="EF90DB890258FF932011F9D9CE231732" author="Korshunova, T. &amp; Sanamyan, N. &amp; Zimina, O. &amp; Fletcher, K. &amp; Martynov, A." box="[805,1077,1571,1598]" journalOrPublisher="ZooKeys" pageId="17" pageNumber="110" pagination="19 - 42" part="630" refId="ref30101" refString="Korshunova, T., Sanamyan, N., Zimina, O., Fletcher, K. &amp; Martynov, A. (2016 a) Two new species and a remarkable record of the genus Dendronotus from the North Pacific and Arctic oceans (Nudibranchia). ZooKeys, 630, 19 - 42." title="Two new species and a remarkable record of the genus Dendronotus from the North Pacific and Arctic oceans (Nudibranchia)" type="journal article" year="2016">Korshunova et al., 2016a</bibRefCitation>
,
<bibRefCitation id="EF90DB890258FF93270BF9D9CE971731" author="Korshunova, T. &amp; Nakano, R. &amp; Fletcher, K. &amp; Sanamyan, N. &amp; Martynov, A." box="[1087,1153,1572,1598]" journalOrPublisher="Venus" pageId="17" pageNumber="110" pagination="1 - 14" part="77" refId="ref29910" refString="Korshunova, T., Nakano, R., Fletcher, K., Sanamyan, N. &amp; Martynov, A. (2019 b) First molecular confirmation of the presence of Dendronotus primorjensis Martynov, Sanamyan &amp; Korshunova, 2015 in Japan and new distributional records of Dendronotus species in the North Pacic (Nudibranchia: Dendronotidae). Venus, 77, 1 - 14." title="First molecular confirmation of the presence of Dendronotus primorjensis Martynov, Sanamyan &amp; Korshunova, 2015 in Japan and new distributional records of Dendronotus species in the North Pacic (Nudibranchia: Dendronotidae)" type="journal article" year="2019">2019b</bibRefCitation>
; for
<taxonomicName id="4C01DDFB0258FF932198F9B5C907176E" box="[684,785,1607,1633]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="17" pageNumber="110" phylum="Mollusca" rank="species" species="kalikal">
<emphasis id="B9757A6A0258FF932198F9B5C907176E" box="[684,785,1607,1633]" italics="true" pageId="17" pageNumber="110">D. kalikal</emphasis>
</taxonomicName>
see fig. 4Q). Therefore, when the original descriptions denticles or ridge-like structures was not indicated for species that we could not check, we suppose the presence of at least one of these structures and indicate that in the diagnoses.
</paragraph>
<paragraph id="8BBEA678025BFF9023ABFF0ECB021396" blockId="18.[129,641,171,1742]" pageId="18" pageNumber="111">
Though a pair of salivary glands indeed inserts into the pharynx on the sides of the oesophagus, the knowledge of these is limited only to a few species (e.g.,
<bibRefCitation id="EF90DB89025BFF902280FEA2C86A1076" author="MacFarland, F. M." box="[436,636,351,377]" journalOrPublisher="Mem. Calif. Acad. Sci." pageId="18" pageNumber="111" pagination="1 - 546" part="6" refId="ref30430" refString="MacFarland, F. M. (1966) Studies of opisthobranchiate mollusks of the Pacific coast of North America. Mem. Calif. Acad. Sci., 6, 1 - 546." title="Studies of opisthobranchiate mollusks of the Pacific coast of North America" type="journal article" year="1966">MacFarland, 1966</bibRefCitation>
;
<bibRefCitation id="EF90DB89025BFF9023B5FE7ECB9B1092" author="Valdes, A. &amp; Bouchet, P." box="[129,397,387,413]" journalOrPublisher="Deep-sea Res. Pt" pageId="18" pageNumber="111" pagination="319 - 327" part="45" refId="ref32934" refString="Valdes, A. &amp; Bouchet, P. (1998) Naked in toxic fluids: a nudibranch mollusc from hydrothermal vents. Deep-sea Res. Pt II, 45, 319 - 327." title="Naked in toxic fluids: a nudibranch mollusc from hydrothermal vents" type="journal article" year="1998">Valdes &amp; Bouchet, 1998</bibRefCitation>
;
<bibRefCitation id="EF90DB89025BFF9022AEFE7EC8621092" author="Pola, M. &amp; Stout, C." box="[410,628,387,413]" journalOrPublisher="Zootaxa" pageId="18" pageNumber="111" pagination="45 - 66" part="1960" refId="ref31659" refString="Pola, M. &amp; Stout, C. (2008) Description of the first two tropical Indo-Pacific species of Dendronotus (Gastropoda: Nudibranchia) with new data of the poorly known species Dendronotus gracilis Baba, 1949. Zootaxa, 1960, 45 - 66." title="Description of the first two tropical Indo-Pacific species of Dendronotus (Gastropoda: Nudibranchia) with new data of the poorly known species Dendronotus gracilis Baba, 1949" type="journal article" year="2008">Pola &amp; Stout, 2008</bibRefCitation>
). The salivary glands are usually not figured in the original descriptions (an exception:
<bibRefCitation id="EF90DB89025BFF9023B5FE13CBA71306" author="Valdes, A. &amp; Bouchet, P." box="[129,433,494,521]" journalOrPublisher="Deep-sea Res. Pt" pageId="18" pageNumber="111" pagination="319 - 327" part="45" refId="ref32934" refString="Valdes, A. &amp; Bouchet, P. (1998) Naked in toxic fluids: a nudibranch mollusc from hydrothermal vents. Deep-sea Res. Pt II, 45, 319 - 327." title="Naked in toxic fluids: a nudibranch mollusc from hydrothermal vents" type="journal article" year="1998">Valdes and Bouchet, 1998</bibRefCitation>
) and were never consistently used for taxonomic diagnostics. We therefore do not include salivary glands in the diagnoses of the species of the genus
<taxonomicName id="4C01DDFB025BFF9023B5FD83CB191397" authorityName="Alder &amp; Hancock" authorityYear="1845" box="[129,271,638,664]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="18" pageNumber="111" phylum="Mollusca" rank="genus">
<emphasis id="B9757A6A025BFF9023B5FD83CB191397" box="[129,271,638,664]" italics="true" pageId="18" pageNumber="111">Dendronotus</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="8BBEA678025BFF9023ABFD5FC9011732" blockId="18.[129,641,171,1742]" lastBlockId="18.[676,1189,171,1597]" pageId="18" pageNumber="111">
The radular patterns within the genus
<taxonomicName id="4C01DDFB025BFF9023B5FD3BCB0613EF" authorityName="Alder &amp; Hancock" authorityYear="1845" box="[129,272,710,736]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="18" pageNumber="111" phylum="Mollusca" rank="genus">
<emphasis id="B9757A6A025BFF9023B5FD3BCB0613EF" box="[129,272,710,736]" italics="true" pageId="18" pageNumber="111">Dendronotus</emphasis>
</taxonomicName>
still offer one of the best diagnostic characters (
<bibRefCitation id="EF90DB89025BFF902261FD17C863120B" author="Korshunova, T. &amp; Martynov, A. &amp; Bakken, T. &amp; Evertsen, J. &amp; Fletcher, K. &amp; Mudianta, I. &amp; Saito, H. &amp; Lundin, K. &amp; Schrodl, M. &amp; Picton, B." box="[341,629,746,772]" journalOrPublisher="ZooKeys" pageId="18" pageNumber="111" pagination="1 - 139" part="717" refId="ref29816" refString="Korshunova, T., Martynov, A., Bakken, T., Evertsen, J., Fletcher, K., Mudianta, I., Saito, H., Lundin, K., Schrodl, M. &amp; Picton, B. (2017 a) Polyphyly of the traditional family Flabellinidae affects a major group of Nudibranchia: aeolidacean taxonomic reassessment with descriptions of several new families, genera, and species (Mollusca, Gastropoda). ZooKeys, 717, 1 - 139." title="Polyphyly of the traditional family Flabellinidae affects a major group of Nudibranchia: aeolidacean taxonomic reassessment with descriptions of several new families, genera, and species (Mollusca, Gastropoda)" type="journal article" year="2017">Korshunova et al., 2017a</bibRefCitation>
). However in several species groups quite similar patterns of the central tooth can be observed. These similarities occur between groups/clades of closely related species (for example between species that closely related to
<taxonomicName id="4C01DDFB025BFF902397FC3FCB2712D4" box="[163,305,961,987]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="18" pageNumber="111" phylum="Mollusca" rank="species" species="frondosus">
<emphasis id="B9757A6A025BFF902397FC3FCB2712D4" box="[163,305,961,987]" italics="true" pageId="18" pageNumber="111">D. frondosus</emphasis>
</taxonomicName>
, e.g.,
<taxonomicName id="4C01DDFB025BFF90224FFC3FC83312D4" box="[379,549,962,987]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="18" pageNumber="111" phylum="Mollusca" rank="species" species="primorjensis">
<emphasis id="B9757A6A025BFF90224FFC3FC83312D4" box="[379,549,962,987]" italics="true" pageId="18" pageNumber="111">D. primorjensis</emphasis>
</taxonomicName>
and
<taxonomicName id="4C01DDFB025BFF90215DFC3FCAF412F0" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="18" pageNumber="111" phylum="Mollusca" rank="species" species="venustus">
<emphasis id="B9757A6A025BFF90215DFC3FCAF412F0" italics="true" pageId="18" pageNumber="111">D. venustus</emphasis>
</taxonomicName>
, between
<taxonomicName id="4C01DDFB025BFF902250FC1BCBC012F0" box="[356,470,997,1023]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="18" pageNumber="111" phylum="Mollusca" rank="species" species="lacteus">
<emphasis id="B9757A6A025BFF902250FC1BCBC012F0" box="[356,470,997,1023]" italics="true" pageId="18" pageNumber="111">D. lacteus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C01DDFB025BFF902129FC1BC86F12F0" authorityName="O'Donoghue" authorityYear="1921" box="[541,633,997,1023]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="18" pageNumber="111" phylum="Mollusca" rank="species" species="rufus">
<emphasis id="B9757A6A025BFF902129FC1BC86F12F0" box="[541,633,997,1023]" italics="true" pageId="18" pageNumber="111">D. rufus</emphasis>
</taxonomicName>
, between
<taxonomicName id="4C01DDFB025BFF9023DBFBF7CB53152C" authorityName="sensu Bergh" authorityYear="1886" box="[239,325,1033,1059]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="18" pageNumber="111" phylum="Mollusca" rank="species" species="dalli">
<emphasis id="B9757A6A025BFF9023DBFBF7CB53152C" box="[239,325,1033,1059]" italics="true" pageId="18" pageNumber="111">D. dalli</emphasis>
</taxonomicName>
and
<taxonomicName id="4C01DDFB025BFF9022BEFBF7C817152C" authorityName="Verrill" authorityYear="1880" box="[394,513,1033,1059]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="18" pageNumber="111" phylum="Mollusca" rank="species" species="elegans">
<emphasis id="B9757A6A025BFF9022BEFBF7C817152C" box="[394,513,1033,1059]" italics="true" pageId="18" pageNumber="111">D. elegans</emphasis>
</taxonomicName>
), but also among apparently more distantly related species suchas
<taxonomicName id="4C01DDFB025BFF902232FBAFCBB91564" box="[262,431,1105,1131]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="18" pageNumber="111" phylum="Mollusca" rank="species" species="kamchaticus">
<emphasis id="B9757A6A025BFF902232FBAFCBB91564" box="[262,431,1105,1131]" italics="true" pageId="18" pageNumber="111">D. kamchaticus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C01DDFB025BFF9022D7FBAFC85C1564" authorityName="Martynov, Fujiwara, Tsuchida, R.Nakano, N.Sanamyan, K.Sanamyan, Fletcher &amp; Korshunova" authorityYear="2020" box="[483,586,1105,1131]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="18" pageNumber="111" phylum="Mollusca" rank="species" species="zakuro">
<emphasis id="B9757A6A025BFF9022D7FBAFC85C1564" box="[483,586,1105,1131]" italics="true" pageId="18" pageNumber="111">D. zakuro</emphasis>
</taxonomicName>
, and between
<taxonomicName id="4C01DDFB025BFF9023DCFB8BCB571580" authorityName="sensu Robilliard" authorityYear="1970" box="[232,321,1141,1167]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="18" pageNumber="111" phylum="Mollusca" rank="species" species="albus">
<emphasis id="B9757A6A025BFF9023DCFB8BCB571580" box="[232,321,1141,1167]" italics="true" pageId="18" pageNumber="111">D. albus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C01DDFB025BFF90224DFB8BC8121580" authorityName="Korshunova, Sanamyan, Zimina, Fletcher, Martynov" authorityYear="2016" box="[377,516,1141,1167]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="18" pageNumber="111" phylum="Mollusca" rank="species" species="robilliardi">
<emphasis id="B9757A6A025BFF90224DFB8BC8121580" box="[377,516,1141,1167]" italics="true" pageId="18" pageNumber="111">D. robilliardi</emphasis>
</taxonomicName>
(see fig. 1). Despite these similarities, in a majority of these cases fine-scale differences in morphology of radula can be detected. At the same time, different species which are grouped in the same subclade (fig. 1) can demonstrate considerably different radular patterns, as for example the clade consisting of
<taxonomicName id="4C01DDFB025BFF9022CAFA8CC86C1485" authorityName="Martynov, Fujiwara, Tsuchida, R. Nakano, N. Sanamyan, K. Sanamyan, Fletcher &amp; Korshunova" authorityYear="2020" box="[510,634,1393,1418]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="18" pageNumber="111" phylum="Mollusca" rank="species" species="jamsteci">
<emphasis id="B9757A6A025BFF9022CAFA8CC86C1485" box="[510,634,1393,1418]" italics="true" pageId="18" pageNumber="111">D. jamsteci</emphasis>
</taxonomicName>
,
<taxonomicName id="4C01DDFB025BFF9023B5FA68CB3214A1" authorityName="MacFarland" authorityYear="1966" box="[129,292,1428,1454]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="18" pageNumber="111" phylum="Mollusca" rank="species" species="subramosus">
<emphasis id="B9757A6A025BFF9023B5FA68CB3214A1" box="[129,292,1428,1454]" italics="true" pageId="18" pageNumber="111">D. subramosus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C01DDFB025BFF90226BFA68CBAF14A1" authorityName="sensu Robilliard" authorityYear="1970" box="[351,441,1428,1454]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="18" pageNumber="111" phylum="Mollusca" rank="species" species="albus">
<emphasis id="B9757A6A025BFF90226BFA68CBAF14A1" box="[351,441,1428,1454]" italics="true" pageId="18" pageNumber="111">D. albus</emphasis>
</taxonomicName>
(
<bibRefCitation id="EF90DB89025BFF9022FDFA69C86A14A1" author="Robilliard, G." box="[457,636,1428,1454]" journalOrPublisher="Veliger" pageId="18" pageNumber="111" pagination="433 - 479" part="12" refId="ref31995" refString="Robilliard, G. (1970) The systematics and some aspects of the ecology of the genus Dendronotus. Veliger, 12, 433 - 479." title="The systematics and some aspects of the ecology of the genus Dendronotus" type="journal article" year="1970">Robilliard, 1970</bibRefCitation>
;
<bibRefCitation id="EF90DB89025BFF9023B5FA45CB4E14DD" author="Pola, M. &amp; Stout, C." box="[129,344,1464,1490]" journalOrPublisher="Zootaxa" pageId="18" pageNumber="111" pagination="45 - 66" part="1960" refId="ref31659" refString="Pola, M. &amp; Stout, C. (2008) Description of the first two tropical Indo-Pacific species of Dendronotus (Gastropoda: Nudibranchia) with new data of the poorly known species Dendronotus gracilis Baba, 1949. Zootaxa, 1960, 45 - 66." title="Description of the first two tropical Indo-Pacific species of Dendronotus (Gastropoda: Nudibranchia) with new data of the poorly known species Dendronotus gracilis Baba, 1949" type="journal article" year="2008">Pola &amp; Stout, 2008</bibRefCitation>
;
<bibRefCitation id="EF90DB89025BFF902250FA45C86D14DD" author="Korshunova, T. &amp; Sanamyan, N. &amp; Zimina, O. &amp; Fletcher, K. &amp; Martynov, A." box="[356,635,1464,1491]" journalOrPublisher="ZooKeys" pageId="18" pageNumber="111" pagination="19 - 42" part="630" refId="ref30101" refString="Korshunova, T., Sanamyan, N., Zimina, O., Fletcher, K. &amp; Martynov, A. (2016 a) Two new species and a remarkable record of the genus Dendronotus from the North Pacific and Arctic oceans (Nudibranchia). ZooKeys, 630, 19 - 42." title="Two new species and a remarkable record of the genus Dendronotus from the North Pacific and Arctic oceans (Nudibranchia)" type="journal article" year="2016">Korshunova et al., 2016a</bibRefCitation>
;
<bibRefCitation id="EF90DB89025BFF9023B5FA21CB6D14F9" author="Martynov, A. &amp; Fujiwara, Y. &amp; Tsuchida, S. &amp; Nakano, R. &amp; Sanamyan, N. &amp; Sanamyan, K. &amp; Fletcher, K. &amp; Korshunova, T." box="[129,379,1500,1526]" journalOrPublisher="Zootaxa" pageId="18" pageNumber="111" pagination="495 - 513" part="4747" refId="ref30619" refString="Martynov, A., Fujiwara, Y., Tsuchida, S., Nakano, R., Sanamyan, N., Sanamyan, K., Fletcher, K. &amp; Korshunova, T. (2020 a) Three new species of the genus Dendronotus from Japan and Russia (Mollusca, Nudibranchia). Zootaxa, 4747, 495 - 513." title="Three new species of the genus Dendronotus from Japan and Russia (Mollusca, Nudibranchia)" type="journal article" year="2020">Martynov et al., 2020a</bibRefCitation>
). Furthermore, the radulae of the morphologically very different
<taxonomicName id="4C01DDFB025BFF90215DF9FCCAF91731" authorityName="Korshunova, Sanamyan, Zimina, Fletcher, Martynov" authorityYear="2016" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="18" pageNumber="111" phylum="Mollusca" rank="species" species="robilliardi">
<emphasis id="B9757A6A025BFF90215DF9FCCAF91731" italics="true" pageId="18" pageNumber="111">D. robilliardi</emphasis>
</taxonomicName>
and
<taxonomicName id="4C01DDFB025BFF902213F9D9CB821731" authorityName="Baba" authorityYear="1949" box="[295,404,1572,1598]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="18" pageNumber="111" phylum="Mollusca" rank="species" species="gracilis">
<emphasis id="B9757A6A025BFF902213F9D9CB821731" box="[295,404,1572,1598]" italics="true" pageId="18" pageNumber="111">D. gracilis</emphasis>
</taxonomicName>
demonstrate similar patterns in the presence of a widened central tooth with numerous small lateral denticles without furrows (
<bibRefCitation id="EF90DB89025BFF902268F972CBF717A5" author="Baba, K." box="[348,481,1679,1706]" journalOrPublisher="Iwanami Shoten, Tokyo" pageId="18" pageNumber="111" refId="ref27988" refString="Baba, K. (1949) Opisthobranchia of Sagami Bay collected by His Majesty The Emperor of Japan. Iwanami Shoten, Tokyo." title="Opisthobranchia of Sagami Bay collected by His Majesty The Emperor of Japan" type="book" year="1949">Baba, 1949</bibRefCitation>
;
<bibRefCitation id="EF90DB89025BFF9022C3F972CB1A17C2" author="Korshunova, T. &amp; Sanamyan, N. &amp; Zimina, O. &amp; Fletcher, K. &amp; Martynov, A." journalOrPublisher="ZooKeys" pageId="18" pageNumber="111" pagination="19 - 42" part="630" refId="ref30101" refString="Korshunova, T., Sanamyan, N., Zimina, O., Fletcher, K. &amp; Martynov, A. (2016 a) Two new species and a remarkable record of the genus Dendronotus from the North Pacific and Arctic oceans (Nudibranchia). ZooKeys, 630, 19 - 42." title="Two new species and a remarkable record of the genus Dendronotus from the North Pacific and Arctic oceans (Nudibranchia)" type="journal article" year="2016">Korshunova et al. 2016a</bibRefCitation>
,
<bibRefCitation id="EF90DB89025BFF90222AF94ECB7517C1" author="Korshunova, T. &amp; Nakano, R. &amp; Fletcher, K. &amp; Sanamyan, N. &amp; Martynov, A." box="[286,355,1715,1742]" journalOrPublisher="Venus" pageId="18" pageNumber="111" pagination="1 - 14" part="77" refId="ref29910" refString="Korshunova, T., Nakano, R., Fletcher, K., Sanamyan, N. &amp; Martynov, A. (2019 b) First molecular confirmation of the presence of Dendronotus primorjensis Martynov, Sanamyan &amp; Korshunova, 2015 in Japan and new distributional records of Dendronotus species in the North Pacic (Nudibranchia: Dendronotidae). Venus, 77, 1 - 14." title="First molecular confirmation of the presence of Dendronotus primorjensis Martynov, Sanamyan &amp; Korshunova, 2015 in Japan and new distributional records of Dendronotus species in the North Pacic (Nudibranchia: Dendronotidae)" type="journal article" year="2019">2019b</bibRefCitation>
). The radula undergoes remarkable transformation during the development from early juvenile to adult. The central tooth in earlier juveniles is low and bears fewdistinct lateral denticles. The lateral teeth in early juveniles are very few in number (13) or absent, showing an indistinct denticulation or they are almost smooth. In individuals approaching the adults stage, the central teeth become elongate-triangular with distinct shoulders, lateral denticles become relatively smaller and numerous or completely reduced. Meanwhile, the lateral teeth increase in number and the denticles become more conspicuous. Since parts of juvenile radula may persist in some adults, this may create a considerable confusion regarding the taxonomic importance of radular characters. For instance,
<taxonomicName id="4C01DDFB025BFF902072FCF3C98C1227" authorityName="sensu Bergh" authorityYear="1886" box="[838,922,782,808]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="18" pageNumber="111" phylum="Mollusca" rank="species" species="dalli">
<emphasis id="B9757A6A025BFF902072FCF3C98C1227" box="[838,922,782,808]" italics="true" pageId="18" pageNumber="111">D. dalli</emphasis>
</taxonomicName>
usually has a smooth central teeth without denticles in adult stage, whereas for adults of
<taxonomicName id="4C01DDFB025BFF9020A0FCABC9F41260" authorityName="sensu Bergh" authorityYear="1886" box="[916,994,853,879]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="18" pageNumber="111" phylum="Mollusca" rank="species" species="dalli">
<emphasis id="B9757A6A025BFF9020A0FCABC9F41260" box="[916,994,853,879]" italics="true" pageId="18" pageNumber="111">D. dalli</emphasis>
</taxonomicName>
also denticles on central teeth were reported, which led to the incorrect synonymyzation of
<taxonomicName id="4C01DDFB025BFF9020C3FC63CE5E12B8" authorityName="sensu Bergh" authorityYear="1886" box="[1015,1096,925,951]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="18" pageNumber="111" phylum="Mollusca" rank="species" species="dalli">
<emphasis id="B9757A6A025BFF9020C3FC63CE5E12B8" box="[1015,1096,925,951]" italics="true" pageId="18" pageNumber="111">D. dalli</emphasis>
</taxonomicName>
with
<taxonomicName id="4C01DDFB025BFF9027B8FC63C90612D4" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="18" pageNumber="111" phylum="Mollusca" rank="species" species="frondosus">
<emphasis id="B9757A6A025BFF9027B8FC63C90612D4" italics="true" pageId="18" pageNumber="111">D. frondosus</emphasis>
</taxonomicName>
(
<bibRefCitation id="EF90DB89025BFF902016FC3CC9AA12D4" author="Odhner, N." box="[802,956,961,987]" journalOrPublisher="Mem. Mus. R. Hist. Nat. Belg." pageId="18" pageNumber="111" pagination="1057 - 1128" part="3" refId="ref31525" refString="Odhner, N. (1936) NudibranchiaDendronotacea - A revision of the system. Mem. Mus. R. Hist. Nat. Belg. (Ser. 2), 3, 1057 - 1128." title="NudibranchiaDendronotacea - A revision of the system" type="journal article" year="1936">Odhner, 1936</bibRefCitation>
;
<bibRefCitation id="EF90DB89025BFF9020FDFC3CCE8E12D3" author="MacFarland, F. M." box="[969,1176,961,988]" journalOrPublisher="Mem. Calif. Acad. Sci." pageId="18" pageNumber="111" pagination="1 - 546" part="6" refId="ref30430" refString="MacFarland, F. M. (1966) Studies of opisthobranchiate mollusks of the Pacific coast of North America. Mem. Calif. Acad. Sci., 6, 1 - 546." title="Studies of opisthobranchiate mollusks of the Pacific coast of North America" type="journal article" year="1966">MacFarland, 1966</bibRefCitation>
). This decision was later changed in favour of the validity of
<taxonomicName id="4C01DDFB025BFF902079FBF7C988152C" authorityName="sensu Bergh" authorityYear="1886" box="[845,926,1033,1059]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="18" pageNumber="111" phylum="Mollusca" rank="species" species="dalli">
<emphasis id="B9757A6A025BFF902079FBF7C988152C" box="[845,926,1033,1059]" italics="true" pageId="18" pageNumber="111">D. dalli</emphasis>
</taxonomicName>
, but uncertanties over the taxonomic application of the putatively variable radula remained (
<bibRefCitation id="EF90DB89025BFF9020D1FBACCE891564" author="Robilliard, G." box="[997,1183,1105,1131]" journalOrPublisher="Veliger" pageId="18" pageNumber="111" pagination="433 - 479" part="12" refId="ref31995" refString="Robilliard, G. (1970) The systematics and some aspects of the ecology of the genus Dendronotus. Veliger, 12, 433 - 479." title="The systematics and some aspects of the ecology of the genus Dendronotus" type="journal article" year="1970">Robilliard, 1970</bibRefCitation>
;
<bibRefCitation id="EF90DB89025BFF902190FB88C9631580" author="Roginskaya, I. S." box="[676,885,1141,1167]" journalOrPublisher="Keys to the Fauna of USSR" pageId="18" pageNumber="111" pagination="155 - 201" part="151" refId="ref32091" refString="Roginskaya, I. S. (1987) Order Nudibranchia Blainville, 1814. In: Molluscs of the White Sea. Keys to the Fauna of USSR, 151, 155 - 201." title="Order Nudibranchia Blainville, 1814" type="journal article" volumeTitle="Molluscs of the White Sea" year="1987">Roginskaya, 1987</bibRefCitation>
). Such “variations” can be overcome by a consistent application of general ontogenetic principles in taxonomy, using the attribution of a particular set of characters to a particular ontogenetic stage (
<bibRefCitation id="EF90DB89025BFF9021C4FAD5C9EE144C" author="Martynov, A. &amp; Ishida, Y. &amp; Irimura, S. &amp; Tajiri, R. &amp; O'Hara, T. &amp; Fujita, T." box="[752,1016,1320,1347]" journalOrPublisher="PLOS ONE" pageId="18" pageNumber="111" pagination="0139463" part="10" refId="ref30687" refString="Martynov, A., Ishida, Y., Irimura, S., Tajiri, R., O'Hara, T. &amp; Fujita, T. (2015 b) When ontogeny matters: a new Japanese species of brittle star illustrates the importance of considering both adult and juvenile characters in taxonomic practice. PLOS ONE, 10, e 0139463." title="When ontogeny matters: a new Japanese species of brittle star illustrates the importance of considering both adult and juvenile characters in taxonomic practice" type="journal article" year="2015">Martynov et al., 2015b</bibRefCitation>
,
<bibRefCitation id="EF90DB89025BFF902731FAD5CE46144C" author="Korshunova, T. &amp; Malmberg, K. &amp; Prkic, J. &amp; Petani, A. &amp; Fletcher, K. &amp; Lundin, K. &amp; Martynov, A." box="[1029,1104,1320,1347]" journalOrPublisher="ZooKeys" pageId="18" pageNumber="111" pagination="15 - 50" part="917" refId="ref29648" refString="Korshunova, T., Malmberg, K., Prkic, J., Petani, A., Fletcher, K., Lundin, K. &amp; Martynov, A. (2020 b) Fine-scale species delimitation: speciation in process and periodic patterns in nudibranch diversity. ZooKeys, 917, 15 - 50." title="Fine-scale species delimitation: speciation in process and periodic patterns in nudibranch diversity" type="journal article" year="2020">2020b</bibRefCitation>
). Thus, the application of a multilevel diagnostic within the genus
<taxonomicName id="4C01DDFB025BFF9020BFFA8DCE0A1485" authorityName="Alder &amp; Hancock" authorityYear="1845" box="[907,1052,1392,1418]" class="Gastropoda" family="Dendronotidae" genus="Dendronotus" kingdom="Animalia" order="Nudibranchia" pageId="18" pageNumber="111" phylum="Mollusca" rank="genus">
<emphasis id="B9757A6A025BFF9020BFFA8DCE0A1485" box="[907,1052,1392,1418]" italics="true" pageId="18" pageNumber="111">Dendronotus</emphasis>
</taxonomicName>
(one that combines morphological characters of various levels and molecular data) instead of a search for immediate and clearly distinctive differences, requires a fine-scale, complex approach.
</paragraph>
</subSubSection>
</treatment>
</document>