treatments-xml/data/33/4A/F7/334AF7B6ED3B18C7274F15F1C4741775.xml
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<mods:title id="823C207A76C85E4E8EBECD2DB059B2E3">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)</mods:title>
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<paragraph id="143B85AE4611F57B4CE0BEAD0B693CFD" pageId="11" pageNumber="12">
<taxonomicName id="311DAA38DDC1BE2F077CFC2A57ECF345" ID-CoL="7PBRR" authority="Miller, 1971" authorityName="Miller" authorityYear="1971" class="Gastropoda" family="Paracoryphellidae" genus="Paracoryphella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Paracoryphella" order="Nudibranchia" pageId="11" pageNumber="12" phylum="Mollusca" rank="genus">
<pageBreakToken id="EF7D971824DAD335FE368015459CF14A" pageId="11" pageNumber="12" start="start">Paracoryphella</pageBreakToken>
Miller, 1971
</taxonomicName>
Figs 2, 7, 8, 9, 10
</paragraph>
</subSubSection>
<subSubSection id="7B7B3A17FBB626F92E259388D8DA3489" pageId="12" pageNumber="13" type="type species">
<paragraph id="63DD32D9BED9A210F4615138EE9A114A" pageId="12" pageNumber="13">
<pageBreakToken id="A40511B469A8A1436BB4A5698BAD11B6" pageId="12" pageNumber="13" start="start">Type</pageBreakToken>
species.
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<paragraph id="ECD5E8F4F20D200C14D4938BD91012E1" pageId="12" pageNumber="13">
<taxonomicName id="D585F374CB6B4079A0A73EB19E0CDB9D" class="Gastropoda" family="Coryphellidae" genus="Coryphella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Coryphella islandica" order="Nudibranchia" pageId="12" pageNumber="13" phylum="Mollusca" rank="species" species="islandica">Coryphella islandica</taxonomicName>
Odhner, 1937
</paragraph>
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<subSubSection id="796AF80610C8CB465662EE1A5B232071" pageId="12" pageNumber="13" type="diagnosis">
<paragraph id="1F52EE8F812B212F9C931BF5B0B7E47C" pageId="12" pageNumber="13">Diagnosis.</paragraph>
<paragraph id="96C4B3A62A6D47765233083B883B481F" pageId="12" pageNumber="13">Body wide. Notal edge present, well-defined, continuous. Cerata not stalked, continuous. Rhinophores smooth to wrinkled, shorter than or similar in size to oral tentacles. Anterior foot corners present. Anus pleuroproctic under the notal edge. Rachidian teeth with strong cusp; lateral denticles not clearly delineated from cusp. Lateral teeth weakly denticulated without attenuated process basally. Reduced additional rows of of small lateral teeth may present. Single distal receptaculum seminis. Long vas deferens without separate granulated prostate. Penis not internal, permanently attached externally.</paragraph>
</subSubSection>
<subSubSection id="5DC2F2D2C86663FE82B7790726176D20" pageId="12" pageNumber="13" type="species included">
<paragraph id="33F978D42DA94F4954A966B8482DEE65" pageId="12" pageNumber="13">Species included.</paragraph>
<paragraph id="8F218DD82EE29E6CBEC5CCFC549AB467" pageId="12" pageNumber="13">
<taxonomicName id="202A7E9958425739147E154E010E0656" class="Gastropoda" family="Paracoryphellidae" genus="Paracoryphella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Paracoryphella ignicrystalla" order="Nudibranchia" pageId="12" pageNumber="13" phylum="Mollusca" rank="species" species="ignicrystalla">Paracoryphella ignicrystalla</taxonomicName>
sp. n. (Fig. 8),
<taxonomicName id="908891D4902CEA11335A41AD8786D7D7" lsidName="P. islandica" pageId="12" pageNumber="13" rank="species" species="islandica">P. islandica</taxonomicName>
(Odhner, 1937) (Fig. 9) (original description in
<bibRefCitation id="BF9926B6D9FA55A6A50C6DBC6325C0CF" author="Odhner, NH" journalOrPublisher="Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening" pageId="81" pageNumber="82" pagination="253 - 257" title="Coryphellaislandica n. sp., a new nudibranchiate mollusc from Iceland." volume="101" year="1937">Odhner 1937</bibRefCitation>
),
<taxonomicName id="DFB7C666998FD3CBF8EECD7638A42BBB" class="Gastropoda" family="Paracoryphellidae" genus="Paracoryphella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Paracoryphella parva" order="Nudibranchia" pageId="12" pageNumber="13" phylum="Mollusca" rank="species" species="parva">Paracoryphella parva</taxonomicName>
(Hadfield, 1963), comb. n. (original description in
<bibRefCitation id="DFAB39B3AE84A1FA827AB7B10622C594" pageId="12" pageNumber="13">Hadfield 1963</bibRefCitation>
) (Fig. 10).
</paragraph>
</subSubSection>
<subSubSection id="E4B9F00E47C14A5B63E37C87BFAF46BA" lastPageId="13" lastPageNumber="14" pageId="12" pageNumber="13" type="remarks">
<paragraph id="62582AE8CAC1FF6809D17E998A7DE0E4" pageId="12" pageNumber="13">Remarks.</paragraph>
<paragraph id="788EB23EA90DDA0BCEAEDF4A5DFBF257" pageId="12" pageNumber="13">
The genus
<taxonomicName id="223904B949E6FD8BD9C2E1538CB1FDDB" class="Gastropoda" family="Paracoryphellidae" genus="Paracoryphella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Paracoryphella" order="Nudibranchia" pageId="12" pageNumber="13" phylum="Mollusca" rank="genus">Paracoryphella</taxonomicName>
and the family
<taxonomicName id="DC2DDECCB29EF9B395B8E64D35EC1FB8" family="Paracoryphellidae" lsidName="" pageId="12" pageNumber="13" rank="family">Paracoryphellidae</taxonomicName>
were initially proposed by
<bibRefCitation id="6A1DB8B89457FAF6D3A8FDC9D706A10C" author="Miller, MC" journalOrPublisher="Zoological Journal of the Linnean Society" pageId="81" pageNumber="82" pagination="311 - 337" title="Aeolid nudibranchs (Gastropoda: Opisthobranchia) of the families Flabellinidae and Eubranchidae from New Zealand waters." url="https://doi.org/10.1111/j.1096-3642.1971.tb00766.x" volume="50" year="1971">Miller (1971)</bibRefCitation>
because of the putative presence of a second asymmetrical row of lateral teeth in the original description of the species &quot;
<taxonomicName id="EF185F1B49C6FBBEFBAED09144F004B8" class="Gastropoda" family="Coryphellidae" genus="Coryphella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Coryphella" order="Nudibranchia" pageId="12" pageNumber="13" phylum="Mollusca" rank="genus">Coryphella</taxonomicName>
&quot;
<taxonomicName id="D7B751EDCBC40AC3F7175E6F7CC6C812" genus="Aeolidacea" lsidName="Aeolidacea islandica" pageId="12" pageNumber="13" rank="species" species="islandica">islandica</taxonomicName>
(see
<bibRefCitation id="D507DFC68925BA9A9B4AA8FAAEC9A8DA" author="Odhner, NH" journalOrPublisher="Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening" pageId="81" pageNumber="82" pagination="253 - 257" title="Coryphellaislandica n. sp., a new nudibranchiate mollusc from Iceland." volume="101" year="1937">Odhner 1937</bibRefCitation>
). However, the latter character is not very evident compared to the true unique feature of the genus
<taxonomicName id="709C7E51173265E0F33633356D1A6069" class="Gastropoda" family="Paracoryphellidae" genus="Paracoryphella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Paracoryphella" order="Nudibranchia" pageId="12" pageNumber="13" phylum="Mollusca" rank="genus">Paracoryphella</taxonomicName>
, a non-retractable, permanently external penis, which is attached directly to the body wall and does not possesses any penial sheath (Figs 8J, 9M, 10C). Importantly, all three known species of this genus invariably possess this feature. This character is unique not only within traditional flabellinids, but also within the majority of
<taxonomicName id="FC6EBFDBDB0FED7F99137B118BB1F0D7" lsidName="Aeolidacea" pageId="12" pageNumber="13" rank="subOrder" subOrder="Aeolidacea">Aeolidacea</taxonomicName>
. Only a single species of the family
<taxonomicName id="9D5AB9CEE31BADC79C690F586BA7EE86" family="Notaeolidiidae" lsidName="" pageId="12" pageNumber="13" rank="family">Notaeolidiidae</taxonomicName>
also has such an external penis (
<bibRefCitation id="D57A2D1769542350D694F132C0080309" author="Waegele, H" journalOrPublisher="Zoologica Scripta" pageId="85" pageNumber="86" pagination="309 - 330" title="Revision of the Antarctic genus Notaeolidia (Gastropoda, Nudibranchia), with a description of a new species." url="https://doi.org/10.1111/j.1463-6409.1990.tb00261.x" volume="19" year="1990">
<normalizedToken id="BB51F98B4E13B9AAD678D7E2E8E8B9C4" originalValue="Wägele">Waegele</normalizedToken>
1990
</bibRefCitation>
). Furthermore, several members of the notaspid family
<taxonomicName id="454D39BBA16E1B5240A39E86B50726B4" family="Pleurobranchidae" lsidName="" pageId="12" pageNumber="13" rank="family">Pleurobranchidae</taxonomicName>
with an internal shell (
<bibRefCitation id="9BB3BC97B0B9DD11AE44E8243FC5DBF6" author="Martynov, AV" journalOrPublisher="Polar Biology" pageId="80" pageNumber="81" pagination="53 - 70" title="The new Arctic side-gilled sea slug genus Boreoberthella (Gastropoda, Opisthobranchia): Pleurobranchoidean systematics and evolution revisited." url="https://doi.org/10.1007/s00300-008-0503-3" volume="32" year="2008">
Martynov and
<normalizedToken id="214A54BE81850660E75A4B0098AAB603" originalValue="Schrödl">Schroedl</normalizedToken>
2008
</bibRefCitation>
) and very basal
<taxonomicName id="A796B1FB89249EE0C12E4F475AAC3DD3" family="Acteonidae" lsidName="" pageId="12" pageNumber="13" rank="family">Acteonidae</taxonomicName>
with an external solid shell also possess an external penis. Therefore, this character within the genus
<taxonomicName id="7937714C9211EF8653FFA1D6B9A5B722" class="Gastropoda" family="Paracoryphellidae" genus="Paracoryphella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Paracoryphella" order="Nudibranchia" pageId="12" pageNumber="13" phylum="Mollusca" rank="genus">Paracoryphella</taxonomicName>
may be either a basal plesiomorphy, or an ontogenetic reversion to the basal plesiomorphy. Molecular data shows that in either case it occurs within one of the most basal clades of the traditional flabellinids. Additionally, analysis of the light microscopy images of the radula of both the type species
<taxonomicName id="A20927311FC775C5159BF6F770EAA6E5" lsidName="P. islandica" pageId="12" pageNumber="13" rank="species" species="islandica">P. islandica</taxonomicName>
and the new species confirm the possible presence of 1-3 very reduced, asymmetrically placed, additional rows of lateral teeth (Figs 8F, 9K). During preparation for the SEM study these apparently reduced additional teeth became fully indistinguishable. Their correspondence to the normal lateral teeth needs to be further investigated.
</paragraph>
<paragraph id="8D6DC2614B35A885CEE1E154DEAC84E2" lastPageId="13" lastPageNumber="14" pageId="12" pageNumber="13">
<taxonomicName id="2521DEABC2E4B855E185E95EFED69F52" class="Gastropoda" family="Coryphellidae" genus="Coryphella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Coryphella parva" order="Nudibranchia" pageId="12" pageNumber="13" phylum="Mollusca" rank="species" species="parva">Coryphella parva</taxonomicName>
, only known from its original description from Swedish waters (
<bibRefCitation id="A800FEEF07AFC21FE66B5131FA4F37D5" pageId="12" pageNumber="13">Hadfield 1963</bibRefCitation>
), was described as having a permanent external penis and therefore is included here in the genus
<taxonomicName id="0D7800E877BC62E8F6C88C494E89BCD7" class="Gastropoda" family="Paracoryphellidae" genus="Paracoryphella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Paracoryphella" order="Nudibranchia" pageId="12" pageNumber="13" phylum="Mollusca" rank="genus">Paracoryphella</taxonomicName>
. Here, for the first time, we have studied type material of
<taxonomicName id="203B52048D39BAA499B25D86D123E1DA" lsidName="C. parva" pageId="12" pageNumber="13" rank="species" species="parva">C. parva</taxonomicName>
from the Natural History Museum of Denmark (NHMD-91476) (Fig. 10) and confirmed that it possesses a non-retractable external penis (Fig. 10C) and other external features that align with the diagnosis of the genus
<taxonomicName id="61890C8782631F64C82458B74E1F9C2C" class="Gastropoda" family="Paracoryphellidae" genus="Paracoryphella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Paracoryphella" order="Nudibranchia" pageId="12" pageNumber="13" phylum="Mollusca" rank="genus">Paracoryphella</taxonomicName>
. The drawings of the radula (Fig. 10D, E) and reproductive systems (Fig. 10F) in the
<pageBreakToken id="16477817D13FBF195CE76917F58F5D72" pageId="13" pageNumber="14" start="start">original</pageBreakToken>
description of
<taxonomicName id="713147DE953DE4E78D42D1EA0D439CDF" lsidName="P. parva" pageId="13" pageNumber="14" rank="species" species="parva">P. parva</taxonomicName>
in
<bibRefCitation id="E28C2B56DC727095510254971C499278" pageId="13" pageNumber="14">Hadfield (1963)</bibRefCitation>
are also very consistent with two other species of the genus
<taxonomicName id="3872B58B6FBE4C3B1467A6FDB7D5B55F" class="Gastropoda" family="Paracoryphellidae" genus="Paracoryphella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Paracoryphella" order="Nudibranchia" pageId="13" pageNumber="14" phylum="Mollusca" rank="genus">Paracoryphella</taxonomicName>
. Remarkably, even though the length of living specimens of
<taxonomicName id="4D43A81141AAFEA4E1CAB966C6A73FA6" lsidName="P. parva" pageId="13" pageNumber="14" rank="species" species="parva">P. parva</taxonomicName>
do not exceed 3.5 mm (fixed not more than 2 mm, Fig. 10A, B), the animals at that size were fully mature and produced egg masses (
<bibRefCitation id="1C3A9EC99BDBBB1CC999C75E9C009896" pageId="13" pageNumber="14">Hadfield 1963</bibRefCitation>
). Because both
<taxonomicName id="1752650E3D9D10CAC55103B9EAB74D7D" lsidName="P. ignicrystalla" pageId="13" pageNumber="14" rank="species" species="ignicrystalla">P. ignicrystalla</taxonomicName>
sp. n. and the type species of the genus
<taxonomicName id="00EE49E79D8361C4A89D1E2A40172158" lsidName="P. islandica" pageId="13" pageNumber="14" rank="species" species="islandica">P. islandica</taxonomicName>
reach mature size in specimens at least three times larger than
<taxonomicName id="CE0B4F8D02363FEBE61D591DD11C7F70" lsidName="P. parva" pageId="13" pageNumber="14" rank="species" species="parva">P. parva</taxonomicName>
, the latter species thus is clearly a separate one and may represent an example of a partial paedomorphosis. According to the molecular phylogenetic analysis, the genus
<taxonomicName id="68BAC0AE49CC88129B84A87E8AED6081" class="Gastropoda" family="Paracoryphellidae" genus="Paracoryphella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Paracoryphella" order="Nudibranchia" pageId="13" pageNumber="14" phylum="Mollusca" rank="genus">Paracoryphella</taxonomicName>
is the sister of the
<taxonomicName id="A49D92119A337A3013BA64FF093AFF4F" class="Gastropoda" family="Paracoryphellidae" genus="Chlamylla" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Chlamylla" order="Nudibranchia" pageId="13" pageNumber="14" phylum="Mollusca" rank="genus">Chlamylla</taxonomicName>
clade (Figs 1, 2, 7). We retain the genus
<taxonomicName id="0DEB20D6E81746CD08ACB2DD4CF7EEB9" class="Gastropoda" family="Paracoryphellidae" genus="Paracoryphella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Paracoryphella" order="Nudibranchia" pageId="13" pageNumber="14" phylum="Mollusca" rank="genus">Paracoryphella</taxonomicName>
as separate because of unique morphological characteristics including a permanent external penis and also the apparent presence of additional rudimentary lateral teeth rows.
</paragraph>
</subSubSection>
</treatment>
</document>