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<mods:title id="19529453FE98A9049A9A101E7B80A3A3">Enigmas from the past: M’Intosh’s (1885) annelidicolous copepods from the voyage of H. M. S. Challenger</mods:title>
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<paragraph id="8BB90A0CFFBFFF800D8FFA40FE01FA12" blockId="24.[151,487,1412,1438]" box="[151,487,1412,1438]" pageId="24" pageNumber="379">
<heading id="D0F1BD60FFBFFF800D8FFA40FE01FA12" bold="true" box="[151,487,1412,1438]" fontSize="11" level="1" pageId="24" pageNumber="379" reason="1">
<taxonomicName id="4C06718FFFBFFF800D8FFA40FE01FA12" authority="Gravier, 1918a" authorityName="Gravier" authorityYear="1918" box="[151,487,1412,1438]" class="Maxillopoda" family="Bradophilidae" genus="Flabellicola" higherTaxonomySource="CoL" kingdom="Animalia" order="Siphonostomatoida" pageId="24" pageNumber="379" phylum="Arthropoda" rank="genus">
<emphasis id="B972D61EFFBFFF800D8FFA40FE01FA12" bold="true" box="[151,487,1412,1438]" pageId="24" pageNumber="379">
<emphasis id="B972D61EFFBFFF800D8FFA40FECEFA12" bold="true" box="[151,296,1412,1438]" italics="true" pageId="24" pageNumber="379">Flabellicola</emphasis>
<bibRefCitation id="EF9777FDFFBFFF800C37FA40FE01FA12" author="Gravier" box="[303,487,1412,1438]" pageId="24" pageNumber="379" refString="Gravier, C. J. (1918 a) Sur un nouveau Copepode (Flabellicola n. g. neapolitana n. sp.) parasite d'un Annelide polychete [Flabelligera diplochaitos (Otto)]. Comptes rendus hebdomadaires des Seances de l'Academie des Sciences, 166, 502 - 505. [published 25 March]" type="journal article" year="1918" yearSuffix="a">Gravier, 1918a</bibRefCitation>
</emphasis>
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<materialsCitation id="3B6E0051FFBFFF810D8FFA0EFF39FD7C" ID-GBIF-Occurrence="1316540606" county="Gulf of Naples. Although" lastPageId="25" lastPageNumber="380" location="The" municipality="Gravier" pageId="24" pageNumber="379" specimenCount="1" typeStatus="holotype">
<location id="8ED95CD7FFBFFF800D8FFA0EFF25FA6F" LSID="urn:lsid:plazi:treatment:03AFBB1AFFBFFF810D8FFA40FF06F96C:8ED95CD7FFBFFF800D8FFA0EFF25FA6F" box="[151,195,1482,1507]" county="Gulf of Naples. Although" municipality="Gravier" name="The" pageId="24" pageNumber="379">The</location>
monotypic genus
<taxonomicName id="4C06718FFFBFFF800CBBFA0EFDCAFA6F" box="[419,556,1482,1507]" class="Maxillopoda" family="Bradophilidae" genus="Flabellicola" higherTaxonomySource="CoL" kingdom="Animalia" order="Siphonostomatoida" pageId="24" pageNumber="379" phylum="Arthropoda" rank="genus">
<emphasis id="B972D61EFFBFFF800CBBFA0EFDCAFA6F" box="[419,556,1482,1507]" italics="true" pageId="24" pageNumber="379">Flabellicola</emphasis>
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was described by
<bibRefCitation id="EF9777FDFFBFFF800E0AFA0EFC2AFA6F" author="Gravier" box="[786,972,1482,1507]" pageId="24" pageNumber="379" refString="Gravier, C. J. (1918 a) Sur un nouveau Copepode (Flabellicola n. g. neapolitana n. sp.) parasite d'un Annelide polychete [Flabelligera diplochaitos (Otto)]. Comptes rendus hebdomadaires des Seances de l'Academie des Sciences, 166, 502 - 505. [published 25 March]" type="journal article" year="1918" yearSuffix="a">
<collectingMunicipality id="6BDD9076FFBFFF800E0AFA0EFC81FA6F" box="[786,871,1482,1507]" pageId="24" pageNumber="379">Gravier</collectingMunicipality>
(1918a)
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from specimens, at different stages of development, parasitic on the flabelligerid polychaete
<taxonomicName id="4C06718FFFBFFF800FEDFA2AFB45F98B" authority="Otto, 1821" authorityName="Otto" authorityYear="1821" box="[757,1187,1518,1543]" class="Polychaeta" family="Flabelligeridae" genus="Flabelligera" kingdom="Animalia" order="Terebellida" pageId="24" pageNumber="379" phylum="Annelida" rank="species" species="diplochaitos">
<emphasis id="B972D61EFFBFFF800FEDFA2AFBF5F98B" box="[757,1043,1518,1543]" italics="true" pageId="24" pageNumber="379">Flabelligera diplochaitos</emphasis>
(
<bibRefCitation id="EF9777FDFFBFFF800939FA2BFB7DF98B" author="Otto" box="[1057,1179,1519,1543]" pageId="24" pageNumber="379" refString="Otto, A. G. (1821) Animalium maritimorum nondum editorum genera duo. Nova Acta physico-medica Academiae caesareae leopoldino-carolinae naturae curiosorum, 10, 617 - 634, plates L - LI." type="journal article" year="1821">Otto, 1821</bibRefCitation>
)
</taxonomicName>
in the
<collectingCounty id="62D87280FFBFFF8009F7FA2AFEE7F9A7" pageId="24" pageNumber="379">Gulf of Naples. Although</collectingCounty>
being abundant at the
<typeStatus id="54BDB4AEFFBFFF800F14F9D7FDDAF9A7" box="[524,572,1555,1579]" pageId="24" pageNumber="379">type</typeStatus>
locality, particularly during April-May when prevalence can be up to 50% (
<bibRefCitation id="EF9777FDFFBFFF800D86F9F2FEA8F9C3" author="Gravier" box="[158,334,1590,1615]" pageId="24" pageNumber="379" refString="Gravier, C. J. (1918 b) Contribution a l'etude d'un Copepode (Flabellicola neapolitana Gravier) parasite d'un Annelide Polychete [Flabelligera (Siphonostoma) diplochaitos (Otto)]. Pubblicazioni della Stazione zoologica di Napoli, 2, 209 - 222. [published 15 October]" type="journal article" year="1918" yearSuffix="b">Gravier 1918b</bibRefCitation>
), the parasite has not been recorded again since its original description. Adult females of
<taxonomicName id="4C06718FFFBFFF800D8FF99EFD82F9FF" authority="Gravier, 1918a" authorityName="Gravier" authorityYear="1918" box="[151,612,1626,1651]" class="Maxillopoda" family="Bradophilidae" genus="Flabellicola" higherTaxonomySource="CoL" kingdom="Animalia" order="Siphonostomatoida" pageId="24" pageNumber="379" phylum="Arthropoda" rank="species" species="neapolitana">
<emphasis id="B972D61EFFBFFF800D8FF99EFE56F9FF" box="[151,432,1626,1651]" italics="true" pageId="24" pageNumber="379">Flabellicola neapolitana</emphasis>
<bibRefCitation id="EF9777FDFFBFFF800CA3F99EFD82F9FF" author="Gravier" box="[443,612,1626,1651]" pageId="24" pageNumber="379" refString="Gravier, C. J. (1918 a) Sur un nouveau Copepode (Flabellicola n. g. neapolitana n. sp.) parasite d'un Annelide polychete [Flabelligera diplochaitos (Otto)]. Comptes rendus hebdomadaires des Seances de l'Academie des Sciences, 166, 502 - 505. [published 25 March]" type="journal article" year="1918" yearSuffix="a">Gravier, 1918a</bibRefCitation>
</taxonomicName>
are mesoparasitic and typically embed themselves in the gills, located dorsally between the two chaetal bundles that form the cephalic cage. Interestingly, similar to the flabelligerid host utilized by
<taxonomicName id="4C06718FFFBFFF800C0CF966FE75F937" box="[276,403,1698,1723]" class="Maxillopoda" family="Bradophilidae" genus="Bradophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Poecilostomatoida" pageId="24" pageNumber="379" phylum="Arthropoda" rank="genus">
<emphasis id="B972D61EFFBFFF800C0CF966FE75F937" box="[276,403,1698,1723]" italics="true" pageId="24" pageNumber="379">Bradophila</emphasis>
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, their presence is betrayed by the paired egg sacs which extend into the cephalic cage of the polychaete. The pyriform ectosoma is about
<quantity id="4CFEA7E9FFBFFF800F89F902FCD2F952" box="[657,820,1734,1759]" metricMagnitude="-4" metricUnit="m" metricValue="3.2499999999999996" metricValueMax="3.5" metricValueMin="3.0" pageId="24" pageNumber="379" unit="mm" value="0.32499999999999996" valueMax="0.35" valueMin="0.3">0.30–0.35 mm</quantity>
in size (measured along its longest axis) and connects via a short neck-like region (stalk) to an internal vesicle (endosoma) which is about twice the length of its external counterpart. Embedded females lack all trace of appendages, digestive tract and external segmentation; however, the endosoma appears to display a transverse constriction dividing it in an anterior and a posterior part (
<bibRefCitation id="EF9777FDFFBFFF80085DF8F6FF07F8E3" author="Gravier" pageId="24" pageNumber="379" refString="Gravier, C. J. (1918 b) Contribution a l'etude d'un Copepode (Flabellicola neapolitana Gravier) parasite d'un Annelide Polychete [Flabelligera (Siphonostoma) diplochaitos (Otto)]. Pubblicazioni della Stazione zoologica di Napoli, 2, 209 - 222. [published 15 October]" type="journal article" year="1918" yearSuffix="b">Gravier 1918b</bibRefCitation>
). The ovaries are located in the anterior part where they connect to paired oviducts that transport the developing eggs via the remainder of the endosoma and connecting stalk to the external genital apertures of the ectosoma. The paired multiseriate egg sacs are very large, attaining
<quantity id="4CFEA7E9FFBFFF800E88F85AFC12F83A" box="[912,1012,1950,1975]" metricMagnitude="-3" metricUnit="m" metricValue="1.06" pageId="24" pageNumber="379" unit="mm" value="1.06">1.06 mm</quantity>
in size (or over three times the length of the ectosoma). It is not uncommon to observe up to five parasites on the gills of a single host but only the larger specimens are ovigerous. Other females at an earlier stage of development are completely embedded in the branchial tissues and appear to be more numerous than those that have a protruding ectosoma.
<bibRefCitation id="EF9777FDFFBEFF8109F0FF78FA7EFF58" author="Gravier" box="[1256,1432,188,213]" pageId="25" pageNumber="380" refString="Gravier, C. J. (1918 a) Sur un nouveau Copepode (Flabellicola n. g. neapolitana n. sp.) parasite d'un Annelide polychete [Flabelligera diplochaitos (Otto)]. Comptes rendus hebdomadaires des Seances de l'Academie des Sciences, 166, 502 - 505. [published 25 March]" type="journal article" year="1918" yearSuffix="a">Gravier (1918a</bibRefCitation>
,
<bibRefCitation id="EF9777FDFFBEFF810D8FFF1BFF38FF74" author="Gravier" box="[151,222,223,248]" pageId="25" pageNumber="380" refString="Gravier, C. J. (1918 b) Contribution a l'etude d'un Copepode (Flabellicola neapolitana Gravier) parasite d'un Annelide Polychete [Flabelligera (Siphonostoma) diplochaitos (Otto)]. Pubblicazioni della Stazione zoologica di Napoli, 2, 209 - 222. [published 15 October]" type="journal article" year="1918" yearSuffix="b">1918b</bibRefCitation>
) failed to observe males but found spermatozoa, presumably stored in a seminal receptacle, in cross-sections of the female’s ectosoma. Contrary to some published claims (
<bibRefCitation id="EF9777FDFFBEFF810E76FEC0FB7EFE91" author="Bresciani" box="[878,1176,260,285]" pageId="25" pageNumber="380" refString="Bresciani, J. &amp; Lutzen, J. (1966) The anatomy of Aphanodomus terebellae (Levinsen) with remarks on the sexuality of the family Xenocoelomidae nov. fam. (Parasitic Copepoda). Bulletin de la Museum national d'Histoire naturelle, Paris, Serie 2, 37, 787 - 806." type="journal article" year="1966">Bresciani &amp; Lützen 1966</bibRefCitation>
: 804;
<bibRefCitation id="EF9777FDFFBEFF8109FBFEC0FA8FFE90" author="Gotto" box="[1251,1385,260,284]" pageId="25" pageNumber="380" refString="Gotto, R. V. (1979) The association of Copepoda with marine invertebrates. Advances in marine Biology, 16, 1 - 109. http: // dx. doi. org / 10.1016 / S 0065 - 2881 (08) 60292 - 8" type="journal article" year="1979">Gotto 1979</bibRefCitation>
: 34) Gravier did not infer that
<taxonomicName id="4C06718FFFBEFF810CCCFEEDFD9DFECC" box="[468,635,295,320]" class="Maxillopoda" family="Bradophilidae" genus="Flabellicola" higherTaxonomySource="CoL" kingdom="Animalia" order="Siphonostomatoida" pageId="25" pageNumber="380" phylum="Arthropoda" rank="species" species="neapolitana">
<emphasis id="B972D61EFFBEFF810CCCFEEDFD9DFECC" box="[468,635,295,320]" italics="true" pageId="25" pageNumber="380">F. neapolitana</emphasis>
</taxonomicName>
was a self-fertilizing hermaphrodite as previously (but incorrectly) proposed for
<taxonomicName id="4C06718FFFBEFF810C2AFE88FBE5FEE8" authority="(Brumpt, 1897) (Caullery &amp; Mesnil 1919)" authorityName="(Brumpt, 1897) (Caullery &amp; Mesnil" authorityYear="1919" box="[306,1027,332,357]" class="Maxillopoda" family="Xenocoelomatidae" genus="Xenocoeloma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Poecilostomatoida" pageId="25" pageNumber="380" phylum="Arthropoda" rank="species" species="alleni">
<emphasis id="B972D61EFFBEFF810C2AFE88FDF2FEE9" box="[306,532,332,357]" italics="true" pageId="25" pageNumber="380">Xenocoeloma alleni</emphasis>
(
<bibRefCitation id="EF9777FDFFBEFF810F3CFE88FD23FEE8" author="Brumpt" box="[548,709,332,356]" pageId="25" pageNumber="380" refString="Brumpt, E. (1897) Sur un Copepode nouveau (Saccopsis Alleni, nova species, parasite de Polycirrus aurantiacus Grube). Comptes rendus hebdomadaires des Seances de l'Academie des Sciences, Paris, 124, 1464 - 1467." type="journal article" year="1897">Brumpt, 1897</bibRefCitation>
) (
<bibRefCitation id="EF9777FDFFBEFF810FC7FE88FC1DFEE8" author="Caullery" box="[735,1019,332,357]" pageId="25" pageNumber="380" refString="Caullery, M. &amp; Mesnil, F. (1919) Xenocoeloma brumpti C. &amp; M., Copepode parasite de Polycirrus arenivorum C. Bulletin biologique de la France et de la Belgique, 53, 161 - 233." type="journal article" year="1919">Caullery &amp; Mesnil 1919</bibRefCitation>
)
</taxonomicName>
but concluded that the sperm must have been introduced by an as yet undiscovered male. Based on a series of histological sections
<bibRefCitation id="EF9777FDFFBEFF8109F1FEABFA7EFE04" author="Gravier" box="[1257,1432,367,392]" pageId="25" pageNumber="380" refString="Gravier, C. J. (1918 a) Sur un nouveau Copepode (Flabellicola n. g. neapolitana n. sp.) parasite d'un Annelide polychete [Flabelligera diplochaitos (Otto)]. Comptes rendus hebdomadaires des Seances de l'Academie des Sciences, 166, 502 - 505. [published 25 March]" type="journal article" year="1918" yearSuffix="a">Gravier (1918a</bibRefCitation>
,
<bibRefCitation id="EF9777FDFFBEFF810D8FFE50FF38FE21" author="Gravier" box="[151,222,404,429]" pageId="25" pageNumber="380" refString="Gravier, C. J. (1918 b) Contribution a l'etude d'un Copepode (Flabellicola neapolitana Gravier) parasite d'un Annelide Polychete [Flabelligera (Siphonostoma) diplochaitos (Otto)]. Pubblicazioni della Stazione zoologica di Napoli, 2, 209 - 222. [published 15 October]" type="journal article" year="1918" yearSuffix="b">1918b</bibRefCitation>
) attempted to reconstruct the female pathway of the life cycle. Although the infective stage is as yet unknown he assumed that it penetrated the host and developed inside the gill tissues. Developing females lack a functional gut and probably absorb nutrients directly through their specialised integument while eggs start developing and gradually fill up the oviducts. At sexual maturity the female penetrates the host integument a second time and protrudes the ectosoma while remaining attached inside with the endosoma. After mating and fertilization, eggs are deposited formings large paired egg sacs and probably remain attached until eclosion takes place. The proposed life cycle is somewhat similar to that of female splanchnotrophids which also penetrate the body wall of their host twice; however, in
<taxonomicName id="4C06718FFFBEFF810F9CFD55FCCFFD24" box="[644,809,655,680]" class="Maxillopoda" family="Bradophilidae" genus="Flabellicola" higherTaxonomySource="CoL" kingdom="Animalia" order="Siphonostomatoida" pageId="25" pageNumber="380" phylum="Arthropoda" rank="species" species="neapolitana">
<emphasis id="B972D61EFFBEFF810F9CFD55FCCFFD24" box="[644,809,655,680]" italics="true" pageId="25" pageNumber="380">F. neapolitana</emphasis>
</taxonomicName>
mating presumably takes place outside the polychaete host while in members of the
<taxonomicName id="4C06718FFFBEFF810CF6FD70FD28FD41" box="[494,718,692,717]" class="Maxillopoda" family="Splanchnotrophidae" kingdom="Animalia" order="Poecilostomatoida" pageId="25" pageNumber="380" phylum="Arthropoda" rank="family">Splanchnotrophidae</taxonomicName>
it occurs inside the viscera of the nudibranch gastropod (
<bibRefCitation id="EF9777FDFFBEFF810847FD70FF35FD7C" author="Huys" pageId="25" pageNumber="380" refString="Huys, R. (2001) Splanchnotrophid systematics: a case of polyphyly and taxonomic myopia. Journal of crustacean Biology, 21, 106 - 156. http: // dx. doi. org / 10.1651 / 0278 - 0372 (2001) 021 [0106: SSACOP] 2.0. CO; 2" type="journal article" year="2001">Huys 2001</bibRefCitation>
).
</materialsCitation>
</paragraph>
</subSubSection>
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<paragraph id="8BB90A0CFFBEFF810DDFFD38FB9FFBB9" blockId="25.[151,1436,151,1760]" pageId="25" pageNumber="380">
<bibRefCitation id="EF9777FDFFBEFF810DDFFD38FE9EFC98" author="Gravier" box="[199,376,764,789]" pageId="25" pageNumber="380" refString="Gravier, C. J. (1918 a) Sur un nouveau Copepode (Flabellicola n. g. neapolitana n. sp.) parasite d'un Annelide polychete [Flabelligera diplochaitos (Otto)]. Comptes rendus hebdomadaires des Seances de l'Academie des Sciences, 166, 502 - 505. [published 25 March]" type="journal article" year="1918" yearSuffix="a">Gravier (1918a</bibRefCitation>
,
<bibRefCitation id="EF9777FDFFBEFF810C91FD38FE29FC99" author="Gravier" box="[393,463,764,789]" pageId="25" pageNumber="380" refString="Gravier, C. J. (1918 b) Contribution a l'etude d'un Copepode (Flabellicola neapolitana Gravier) parasite d'un Annelide Polychete [Flabelligera (Siphonostoma) diplochaitos (Otto)]. Pubblicazioni della Stazione zoologica di Napoli, 2, 209 - 222. [published 15 October]" type="journal article" year="1918" yearSuffix="b">1918b</bibRefCitation>
) compared
<taxonomicName id="4C06718FFFBEFF810F46FD38FD01FC99" box="[606,743,764,789]" class="Maxillopoda" family="Bradophilidae" genus="Flabellicola" higherTaxonomySource="CoL" kingdom="Animalia" order="Siphonostomatoida" pageId="25" pageNumber="380" phylum="Arthropoda" rank="genus">
<emphasis id="B972D61EFFBEFF810F46FD38FD01FC99" box="[606,743,764,789]" italics="true" pageId="25" pageNumber="380">Flabellicola</emphasis>
</taxonomicName>
with other genera that were known to utilize flabelligeran polychaetes as hosts, claiming that it was probably closest to
<emphasis id="B972D61EFFBEFF810E69FCDBFBF6FCB4" box="[881,1040,799,824]" italics="true" pageId="25" pageNumber="380">Trophoniphila</emphasis>
. Although he suggested a certain affinity with the
<taxonomicName id="4C06718FFFBEFF810C44FC80FDE2FCD1" box="[348,516,836,861]" class="Maxillopoda" family="Herpyllobiidae" kingdom="Animalia" order="Poecilostomatoida" pageId="25" pageNumber="380" phylum="Arthropoda" rank="family">Herpyllobiidae</taxonomicName>
the genus remained unplaced (
<emphasis id="B972D61EFFBEFF810E6AFC80FC68FCD0" box="[882,910,836,860]" italics="true" pageId="25" pageNumber="380">cf.</emphasis>
<bibRefCitation id="EF9777FDFFBEFF810E8FFC80FBFDFCD0" author="Gotto" box="[919,1051,836,860]" pageId="25" pageNumber="380" refString="Gotto, R. V. (1979) The association of Copepoda with marine invertebrates. Advances in marine Biology, 16, 1 - 109. http: // dx. doi. org / 10.1016 / S 0065 - 2881 (08) 60292 - 8" type="journal article" year="1979">Gotto 1979</bibRefCitation>
) until
<bibRefCitation id="EF9777FDFFBEFF810973FC80FA7AFCD0" author="Boxshall" box="[1131,1436,836,861]" pageId="25" pageNumber="380" refString="Boxshall, G. A. &amp; Halsey, H. S. (2004) An Introduction to Copepod Diversity. The Ray Society, London, xv + 966 pp." type="book" year="2004">Boxshall &amp; Halsey (2004)</bibRefCitation>
treated it as a
<emphasis id="B972D61EFFBEFF810C28FCADFDE0FC0C" box="[304,518,871,896]" italics="true" pageId="25" pageNumber="380">genus inquirendum</emphasis>
in the
<taxonomicName id="4C06718FFFBEFF810F49FCA3FD39FC0C" box="[593,735,871,896]" class="Maxillopoda" family="Saccopsidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Poecilostomatoida" pageId="25" pageNumber="380" phylum="Arthropoda" rank="family">Saccopsidae</taxonomicName>
. The complete absence of appendages on the female ectosoma, the size and shape of the egg sacs and its utilization of a flabelligerid polychaete as host suggest a possible relationship with the
<taxonomicName id="4C06718FFFBEFF810C92FC6BFDCFFC44" box="[394,553,943,968]" class="Maxillopoda" family="Bradophilidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Poecilostomatoida" pageId="25" pageNumber="380" phylum="Arthropoda" rank="family">Bradophilidae</taxonomicName>
rather than with the
<taxonomicName id="4C06718FFFBEFF810E02FC6BFC42FC44" box="[794,932,943,968]" class="Maxillopoda" family="Saccopsidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Poecilostomatoida" pageId="25" pageNumber="380" phylum="Arthropoda" rank="family">Saccopsidae</taxonomicName>
which typically have three pairs of reduced cephalic appendages, much smaller, sausage-shaped or straight egg sacs and utilize ampharetids, terebellids and trichobranchids as hosts (
<bibRefCitation id="EF9777FDFFBEFF810CCCFC3CFDB4FB9C" author="Sars" box="[468,594,1016,1040]" pageId="25" pageNumber="380" refString="Sars, M. (1870) VII. Bidrag til Kundskab om Christianiafjordens Fauna. II. Crustacea. Beskrivelse af nye, paa annelider snyltende Copepodeformer. Nyt Magazin for Naturvidenskaberne, 17 (2), 113 - 160, plates 8 - 10." type="journal article" year="1870">Sars 1870</bibRefCitation>
;
<bibRefCitation id="EF9777FDFFBEFF810F71FC33FCFBFB9C" author="Levinsen" box="[617,797,1015,1040]" pageId="25" pageNumber="380" refString="Levinsen, G. M. R. (1878) Om nogle parasitiske Krebsdyr, der snylte hos Annelider. Videnskabelige Meddelelser fra den naturhistoriske Forening i Kjobenhavn, 1877 - 1878, 351 - 380, plate 6." type="journal volume" year="1878">Levinsen 1878</bibRefCitation>
; M’Intosh
<bibRefCitation id="EF9777FDFFBEFF810EB6FC33FC0FFB9C" author="M'Intosh" box="[942,1001,1015,1040]" pageId="25" pageNumber="380" refString="M'Intosh, W. C. (1885) Report on Annelida Polychaeta collected by H. M. S. Challenger during the years 1873 - 1876. Report of the scientific Results of the Voyage of H. M. S. Challenger 1873 - 76, Zoology, 12, i - xxxvi, 1 - 554, plates I - LV, I A - XXXIX A." type="journal article" year="1885">1885</bibRefCitation>
;
<bibRefCitation id="EF9777FDFFBEFF810EE7FC33FADBFB9C" author="Bresciani" box="[1023,1341,1015,1040]" pageId="25" pageNumber="380" refString="Bresciani, J. &amp; Lutzen, J. (1961) The anatomy of a parasitic copepod, Saccopsis steenstrupi n. sp. Crustaceana, 3, 9 - 23. http: // dx. doi. org / 10.1163 / 156854061 X 00491" type="journal article" year="1961">Bresciani &amp; Lützen 1961</bibRefCitation>
,
<bibRefCitation id="EF9777FDFFBEFF81084CFC33FA76FB9C" author="Bresciani" box="[1364,1424,1015,1040]" pageId="25" pageNumber="380" refString="Bresciani, J. &amp; Lutzen, J. (1975) Melinnacheres ergasiloides M. Sars, a parasitic copepod of the polychaete Melinna cristata, with notes on multiple infections caused by annelidicolous copepods. Ophelia, 13, 31 - 42. http: // dx. doi. org / 10.1080 / 00785326.1974.10430590" type="journal article" year="1975">1975</bibRefCitation>
). Consequently,
<taxonomicName id="4C06718FFFBEFF810C25FBD8FE20FBB9" box="[317,454,1052,1077]" class="Maxillopoda" family="Bradophilidae" genus="Flabellicola" higherTaxonomySource="CoL" kingdom="Animalia" order="Siphonostomatoida" pageId="25" pageNumber="380" phylum="Arthropoda" rank="genus">
<emphasis id="B972D61EFFBEFF810C25FBD8FE20FBB9" box="[317,454,1052,1077]" italics="true" pageId="25" pageNumber="380">Flabellicola</emphasis>
</taxonomicName>
is here treated as a
<emphasis id="B972D61EFFBEFF810FBBFBD9FC60FBB9" box="[675,902,1052,1077]" italics="true" pageId="25" pageNumber="380">genus incertae sedis</emphasis>
in the
<taxonomicName id="4C06718FFFBEFF810ECAFBD8FB93FBB9" box="[978,1141,1052,1077]" class="Maxillopoda" family="Bradophilidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Poecilostomatoida" pageId="25" pageNumber="380" phylum="Arthropoda" rank="family">Bradophilidae</taxonomicName>
.
</paragraph>
<paragraph id="8BB90A0CFFBEFF810DDFFBFBFD65FB64" blockId="25.[151,1436,151,1760]" pageId="25" pageNumber="380">
The positioning of the ovaries in the endosoma of
<taxonomicName id="4C06718FFFBEFF810E30FBFBFC57FBD4" box="[808,945,1087,1112]" class="Maxillopoda" family="Bradophilidae" genus="Flabellicola" higherTaxonomySource="CoL" kingdom="Animalia" order="Siphonostomatoida" pageId="25" pageNumber="380" phylum="Arthropoda" rank="genus">
<emphasis id="B972D61EFFBEFF810E30FBFBFC57FBD4" box="[808,945,1087,1112]" italics="true" pageId="25" pageNumber="380">Flabellicola</emphasis>
</taxonomicName>
rather than in its ectosoma is unusual in mesoparasitic copepods and has so far only been observed in the recently discovered
<taxonomicName id="4C06718FFFBEFF810952FBA0FEEEFB2C" authority="Boxshall et al. 2015" authorityName="Boxshall et al." authorityYear="2015" class="Maxillopoda" family="Jasmineiricolidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Poecilostomatoida" pageId="25" pageNumber="380" phylum="Arthropoda" rank="family">
Jasmineiricolidae (
<bibRefCitation id="EF9777FDFFBEFF810804FBA0FEE6FB2C" author="Boxshall" pageId="25" pageNumber="380" refString="Boxshall, G. A., O'Reilly, M., Sikorski, A. &amp; Summerfield, R. (2015) A new genus and family of copepods (Crustacea: Copepoda) parasitic on polychaetes of the genus Jasmineira Langerhans, 1880 (family Sabellidae) in the northeastern Atlantic. Zootaxa, 4018 (3), 426 - 436. http: // dx. doi. org / 10.11646 / zootaxa. 4018.3.6" type="journal article" year="2015">
Boxshall
<emphasis id="B972D61EFFBEFF810890FBA1FF52FB2C" italics="true" pageId="25" pageNumber="380">et al.</emphasis>
2015
</bibRefCitation>
)
</taxonomicName>
. Whether this character is also diagnostic for the
<taxonomicName id="4C06718FFFBEFF810E63FB43FBFCFB2C" box="[891,1050,1159,1184]" class="Maxillopoda" family="Bradophilidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Poecilostomatoida" pageId="25" pageNumber="380" phylum="Arthropoda" rank="family">Bradophilidae</taxonomicName>
remains to be confirmed since
<bibRefCitation id="EF9777FDFFBEFF810D8FFB68FE66FB49" author="Marchenkov" box="[151,384,1196,1221]" pageId="25" pageNumber="380" refString="Marchenkov, A. V. (1999 b) Symbiotic and parasitic Crustacea of benthic marine invertebrates from the Arctic Seas of Russia. Synopsis of Ph. D. dissertation, Saint Petersburg, 23 pp." type="book" year="1999" yearSuffix="b">Marchenkov (1999b</bibRefCitation>
,
<bibRefCitation id="EF9777FDFFBEFF810C95FB68FE21FB48" author="Marchenkov" box="[397,455,1196,1220]" pageId="25" pageNumber="380" refString="Marchenkov, A. V. (2002) Bradophilidae fam. nov. - the new family of mesoparasitic copepod collected from the polychaete Brada villosa from the White Sea. Parazitologiya, 36, 514 - 516. [in Russian with English summary]" type="journal article" year="2002">2002</bibRefCitation>
) did not comment on the position of the ovaries in
<taxonomicName id="4C06718FFFBEFF810904FB68FB7AFB49" box="[1052,1180,1196,1221]" class="Maxillopoda" family="Bradophilidae" genus="Bradophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Poecilostomatoida" pageId="25" pageNumber="380" phylum="Arthropoda" rank="genus">
<emphasis id="B972D61EFFBEFF810904FB68FB7AFB49" box="[1052,1180,1196,1221]" italics="true" pageId="25" pageNumber="380">Bradophila</emphasis>
</taxonomicName>
and no information is available on the condition in
<emphasis id="B972D61EFFBEFF810CC4FB0BFD9DFB64" box="[476,635,1231,1256]" italics="true" pageId="25" pageNumber="380">Trophoniphila</emphasis>
.
</paragraph>
<paragraph id="8BB90A0CFFBEFF810DDFFB30FF06F96C" blockId="25.[151,1436,151,1760]" pageId="25" pageNumber="380">
Flabelligerid polychaetes are rarely reported as hosts for parasitic copepods. The most important diagnostic feature of flabelligerids is their ability to retract the anterior end, including the pro- and peristomium with associated palps and gills, and some of the anterior chaetigerous segments into the body, forming an introvert (
<bibRefCitation id="EF9777FDFFBEFF810D86FA9BFE7CFAF4" author="Filippova" box="[158,410,1375,1400]" pageId="25" pageNumber="380" refString="Filippova, A. V., Tzetlin, A. B. &amp; Purschke, G. (2003) Morphology and ultrastructure of the anterior end of Diplocirrus longisetosus Marenzeller, 1890 (Flabelligeridae, Polychaeta, Annelida). In: Sigvaldadottir, E., Mackie, A. S. Y., Helgason, G. V., Reish, D. J., Svavarsson, J., Steingrimsson, S. A. &amp; Gudmundsson, G. (Eds.), Advances in Polychaete Research. Hydrobiologia, 496, 215 - 223. http: // dx. doi. org / 10.1023 / A: 1026148900074" type="journal article" year="2003">
Filippova
<emphasis id="B972D61EFFBEFF810C0EFAA5FEB5FAF4" box="[278,339,1375,1400]" italics="true" pageId="25" pageNumber="380">et al.</emphasis>
2003
</bibRefCitation>
). This confined microhabitat is the preferred attachment site for the three mesoparasitic members of the
<taxonomicName id="4C06718FFFBEFF810C53FA40FE08FA11" box="[331,494,1412,1437]" class="Maxillopoda" family="Bradophilidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Poecilostomatoida" pageId="25" pageNumber="380" phylum="Arthropoda" rank="family">Bradophilidae</taxonomicName>
. The only other copepod that is known to utilize a flabelligerid host is
<taxonomicName id="4C06718FFFBEFF8109E7FA40FDC7FA4C" authority="Bresciani &amp; Lutzen, 1962" authorityName="Bresciani &amp; Lutzen" authorityYear="1962" class="Maxillopoda" family="Nereicolidae" genus="Flabelliphilus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Poecilostomatoida" pageId="25" pageNumber="380" phylum="Arthropoda" rank="species" species="inersus">
<emphasis id="B972D61EFFBEFF8109E7FA40FF0EFA4C" italics="true" pageId="25" pageNumber="380">Flabelliphilus inersus</emphasis>
Bresciani &amp; Lützen, 1962
</taxonomicName>
, which was described from off the Swedish west coast (Bresciani &amp; Lützen 1962) but has no bradophilid affinities and occupies a different niche on the host. The species inhabits the gelatinous coat of
<taxonomicName id="4C06718FFFBEFF810C6BFA2BFCE0F984" authority="M. Sars, 1829" authorityName="M. Sars" authorityYear="1829" box="[371,774,1519,1544]" class="Polychaeta" family="Flabelligeridae" genus="Flabelligera" kingdom="Animalia" order="Terebellida" pageId="25" pageNumber="380" phylum="Annelida" rank="species" species="affinis">
<emphasis id="B972D61EFFBEFF810C6BFA2BFDB7F984" box="[371,593,1519,1544]" italics="true" pageId="25" pageNumber="380">Flabelligera affinis</emphasis>
<bibRefCitation id="EF9777FDFFBEFF810F44FA35FCE0F984" author="Sars" box="[604,774,1520,1544]" pageId="25" pageNumber="380" refString="Sars, M. (1829) Bidrag til Soedyrenes Naturhistorie. Vol. 1. Chr. Dahl, Bergen, 60 pp., 6 plates." type="book" year="1829">M. Sars, 1829</bibRefCitation>
</taxonomicName>
, is known from males only and generally considered of doubtful systematic position due to the incomplete description.
<bibRefCitation id="EF9777FDFFBEFF810E98F9D0FBF3F9A1" author="Stock" box="[896,1045,1556,1581]" pageId="25" pageNumber="380" refString="Stock, J. H. (1968) Vectoriella marinovi, un Copepode nouveau, parasite d'une Annelide Polychete pontique. Crustaceana, supplement 1, 186 - 192." type="journal article" year="1968">Stock (1968)</bibRefCitation>
listed
<taxonomicName id="4C06718FFFBEFF810971F9D0FEC9F9DC" authority="Bresciani &amp; Lutzen, 1962" authorityName="Bresciani &amp; Lutzen" authorityYear="1962" class="Maxillopoda" family="Nereicolidae" genus="Flabelliphilus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Poecilostomatoida" pageId="25" pageNumber="380" phylum="Arthropoda" rank="genus">
<emphasis id="B972D61EFFBEFF810971F9D0FAE0F9A1" box="[1129,1286,1556,1581]" italics="true" pageId="25" pageNumber="380">Flabelliphilus</emphasis>
Bresciani &amp; Lützen, 1962
</taxonomicName>
as a
<emphasis id="B972D61EFFBEFF810C6CF9FDFDA9F9DC" box="[372,591,1591,1616]" italics="true" pageId="25" pageNumber="380">genus inquirendum</emphasis>
in his “nereicolid series” but
<bibRefCitation id="EF9777FDFFBEFF810EAEF9F3FB85F9DC" author="Laubier" box="[950,1123,1591,1616]" pageId="25" pageNumber="380" refString="Laubier, L. (1978) Ophelicola drachi gen. sp. n., un nouveau Copepode Cyclopoide abyssal ectoparasite d'Annelides Polychetes Opheliidae. Archives de Zoologie experimentale et generale, 119, 39 - 50." type="journal article" year="1978">Laubier (1978)</bibRefCitation>
regarded a position in the Nereicoliform Group unlikely.
<bibRefCitation id="EF9777FDFFBEFF810CEEF998FD55F9F9" author="Bresciani" box="[502,691,1628,1653]" pageId="25" pageNumber="380" refString="Bresciani, J. (1964) Sigecheres brittae gen. et sp. nov., a parasitic copepod from the polychaete Sige fusigera Malmgren. Ophelia, 1, 295 - 301. http: // dx. doi. org / 10.1080 / 00785326.1964.10416284" type="journal article" year="1964">Bresciani (1964)</bibRefCitation>
had previously considered it a possible candidate for inclusion in the
<taxonomicName id="4C06718FFFBEFF810DDAF9BBFEB4F914" box="[194,338,1663,1688]" class="Maxillopoda" family="Clausidiidae" kingdom="Animalia" order="Poecilostomatoida" pageId="25" pageNumber="380" phylum="Arthropoda" rank="family">Clausidiidae</taxonomicName>
.
<bibRefCitation id="EF9777FDFFBEFF810C46F9BBFD6DF914" author="Boxshall" box="[350,651,1663,1688]" pageId="25" pageNumber="380" refString="Boxshall, G. A. &amp; Halsey, H. S. (2004) An Introduction to Copepod Diversity. The Ray Society, London, xv + 966 pp." type="book" year="2004">Boxshall &amp; Halsey (2004)</bibRefCitation>
ranked it as a
<emphasis id="B972D61EFFBEFF810E2FF945FBE9F914" box="[823,1039,1663,1688]" italics="true" pageId="25" pageNumber="380">genus inquirendum</emphasis>
based on the clausiid-like antenna and mandible while subsequent workers cited it as a valid genus of this family (
<bibRefCitation id="EF9777FDFFBEFF81093CF960FB12F930" author="Cheng" box="[1060,1268,1700,1725]" pageId="25" pageNumber="380" refString="Cheng, Y. - r., Ho, J. - s. &amp; Dai, C. - F. (2014) Description of Indoclausia helioporae sp. nov. (Copepoda, Clausiidae) from blue coral, associated with a number of polychaetes. Crustaceana, 87, 627 - 633. http: // dx. doi. org / 10.1163 / 15685403 - 00003309" type="journal article" year="2014">
Cheng
<emphasis id="B972D61EFFBEFF81096EF961FB56F931" box="[1142,1200,1700,1725]" italics="true" pageId="25" pageNumber="380">et al.</emphasis>
2014
</bibRefCitation>
;
<bibRefCitation id="EF9777FDFFBEFF810818F960FF32F96C" author="Conradi" pageId="25" pageNumber="380" refString="Conradi, M., Bandera, M. E., Marin, I. &amp; Martin, D. (2015) Polychaete-parasitizing copepods from the deep-sea Kuril - Kamchatka Trench (Pacific Ocean), with the description of a new Ophelicola species and comments on the currently known annelidicolous copepods. Deep-Sea Research Part II: Topical Studies in Oceanography, 111, 147 - 165. http: // dx. doi. org / 10.1016 / j. dsr 2.2014.08.018" type="journal article" year="2015">
Conradi
<emphasis id="B972D61EFFBEFF81087BF961FA7AF931" box="[1379,1436,1700,1725]" italics="true" pageId="25" pageNumber="380">et al.</emphasis>
2015
</bibRefCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>
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