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<mods:title id="E4C3B2531FDC175C6AC98A5F97170943">New records of bopyrid parasites (Crustacea, Isopoda, Epicaridea) of pea crabs (Crustacea, Decapoda, Pinnotheridae) with descriptions of two new species of Rhopalione Pérez, 1921 and a review of the genus</mods:title>
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<mods:namePart id="9459A005EB61AC0D2A63A5BB7C88C504">Boyko, Christopher B.</mods:namePart>
<mods:affiliation id="98D22629F8956D177CDA45AEB12B3A35">Department of Biology, Hofstra University, 1000 Hempstead Turnpike, Hempstead, NY 11549 (United States); and Division of Invertebrate Zoology, American Museum of Natural History, 200 Central Park West, New York, NY, 10024 (United States) cboyko @ amnh. org (corresponding author)</mods:affiliation>
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<mods:namePart id="A3FB508EDA6B827D80FA6F35F4C68AC7">Williams, Jason D.</mods:namePart>
<mods:affiliation id="5FDB853FA24E95ED0A391603413F070E">Department of Biology, Hofstra University, 1000 Hempstead Turnpike, Hempstead, NY 11549 (United States) jason. d. williams @ hofstra. edu</mods:affiliation>
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Genus
<taxonomicName id="4C1953BFFFB3AB56FBE2FF5FE351FFC4" ID-CoL="7PXNJ" authority="Perez, 1920" authorityName="Perez" authorityYear="1920" box="[1048,1296,215,241]" class="Malacostraca" family="Bopyridae" genus="Rhopalione" kingdom="Animalia" order="Isopoda" pageId="9" pageNumber="156" phylum="Arthropoda" rank="genus">
<emphasis id="B96DF42EFFB3AB56FBE2FF5FE2CAFFC4" box="[1048,1163,215,241]" italics="true" pageId="9" pageNumber="156">Rhopalione</emphasis>
Pérez, 1920
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. —
<taxonomicName id="4C1953BFFFB3AB56FC1BFE98E374FE1D" authority="Perez, 1920" authorityName="Perez" authorityYear="1920" box="[993,1333,272,296]" class="Malacostraca" family="Bopyridae" genus="Rhopalione" kingdom="Animalia" order="Isopoda" pageId="9" pageNumber="156" phylum="Arthropoda" rank="species" species="uromyzon">
<emphasis id="B96DF42EFFB3AB56FC1BFE98E2F5FE1D" box="[993,1204,272,296]" italics="true" pageId="9" pageNumber="156">Rhopalione uromyzon</emphasis>
Pérez, 1920
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, by original designation.
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The occurrence of female bopyrid isopods under the pleons of brachyuran crabs is apparently restricted to members of the genera
<taxonomicName id="4C1953BFFFB3AB56FC59FE5EE257FEC5" authorityName="Perez" authorityYear="1920" box="[931,1046,470,496]" class="Malacostraca" family="Bopyridae" genus="Rhopalione" kingdom="Animalia" order="Isopoda" pageId="9" pageNumber="156" phylum="Arthropoda" rank="genus">
<emphasis id="B96DF42EFFB3AB56FC59FE5EE257FEC5" box="[931,1046,470,496]" italics="true" pageId="9" pageNumber="156">Rhopalione</emphasis>
</taxonomicName>
and
<taxonomicName id="4C1953BFFFB3AB56FBB6FE5EE528FD25" authority="Boyko &amp; van der Meij, 2018" authorityName="Boyko &amp; van der Meij" authorityYear="2018" class="Malacostraca" family="Bopyridae" genus="Spathione" kingdom="Animalia" order="Isopoda" pageId="9" pageNumber="156" phylum="Arthropoda" rank="genus">
<emphasis id="B96DF42EFFB3AB56FBB6FE5EE2F3FEC5" box="[1100,1202,470,496]" italics="true" pageId="9" pageNumber="156">Spathione</emphasis>
<bibRefCitation id="EF8855CDFFB3AB56FB43FE5EE528FD25" author="BOYKO C. B. &amp; VAN DER MEIJ S. E. T." pageId="9" pageNumber="156" pagination="304 - 329" refId="ref20017" refString="BOYKO C. B. &amp; VAN DER MEIJ S. E. T. 2018. - A trifecta of Swiftian symbioses: stony corals, gall crabs and their parasites (Scleractinia; Brachyura: Cryptochiridae; Isopoda: Epicaridea and Cirripedia: Rhizocephala). Zoological Journal of the Linnean Society 184: 304 - 329. https: // doi. org / 10.1093 / zoolinnean / zlx 115" type="journal article" year="2018">Boyko &amp; van der Meij, 2018</bibRefCitation>
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. All other bopyrids that occur abdominally are in subfamilies other than Pseudioninae and occur on anomurans and caridean shrimps but never on brachyurans. This unique site of parasitization might correlate with the morphology of the host carapace. All species of
<taxonomicName id="4C1953BFFFB3AB56FB7FFDFEE2B9FDA5" authorityName="Perez" authorityYear="1920" box="[1157,1272,630,656]" class="Malacostraca" family="Bopyridae" genus="Rhopalione" kingdom="Animalia" order="Isopoda" pageId="9" pageNumber="156" phylum="Arthropoda" rank="genus">
<emphasis id="B96DF42EFFB3AB56FB7FFDFEE2B9FDA5" box="[1157,1272,630,656]" italics="true" pageId="9" pageNumber="156">Rhopalione</emphasis>
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parasitize genera of pinnotherid crabs (Pinnotherinae) in which, for females at least, the carapace is very soft and weakly calcified. This is in marked contrast to the carapace of other brachyurans infested by bopyrids, which is well calcified in both sexes and offers more protection to the parasite; this is true even in the gall crabs (
<taxonomicName id="4C1953BFFFB3AB56FC8BFCBDE2FAFC65" authority="Paulson, 1875" authorityName="Paulson" authorityYear="1875" box="[881,1211,821,848]" class="Malacostraca" family="Cryptochiridae" kingdom="Animalia" order="Decapoda" pageId="9" pageNumber="156" phylum="Arthropoda" rank="family">Cryptochiridae Paulson, 1875</taxonomicName>
), which are parasitized by species of
<taxonomicName id="4C1953BFFFB3AB56FC3FFCDDE26EFC5A" authorityName="Boyko &amp; van der Meij" authorityYear="2018" box="[965,1071,853,879]" class="Malacostraca" family="Bopyridae" genus="Spathione" kingdom="Animalia" order="Isopoda" pageId="9" pageNumber="156" phylum="Arthropoda" rank="genus">
<emphasis id="B96DF42EFFB3AB56FC3FFCDDE26EFC5A" box="[965,1071,853,879]" italics="true" pageId="9" pageNumber="156">Spathione</emphasis>
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. The adaptation to an abdominal mode of infestation by species of
<taxonomicName id="4C1953BFFFB3AB56FB60FCFDE34CFCBA" authorityName="Perez" authorityYear="1920" box="[1178,1293,885,911]" class="Malacostraca" family="Bopyridae" genus="Rhopalione" kingdom="Animalia" order="Isopoda" pageId="9" pageNumber="156" phylum="Arthropoda" rank="genus">
<emphasis id="B96DF42EFFB3AB56FB60FCFDE34CFCBA" box="[1178,1293,885,911]" italics="true" pageId="9" pageNumber="156">Rhopalione</emphasis>
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might have allowed for maximal protection by the host, which is better obtained in an abdominal position; alternatively,
<taxonomicName id="4C1953BFFFB3AB56FAC4FC3DE3F0FCFA" authorityName="Perez" authorityYear="1920" box="[1342,1457,949,975]" class="Malacostraca" family="Bopyridae" genus="Rhopalione" kingdom="Animalia" order="Isopoda" pageId="9" pageNumber="156" phylum="Arthropoda" rank="genus">
<emphasis id="B96DF42EFFB3AB56FAC4FC3DE3F0FCFA" box="[1342,1457,949,975]" italics="true" pageId="9" pageNumber="156">Rhopalione</emphasis>
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is a relatively large bopyrid and the space afforded under the pleon is much larger than the branchial chamber (see further remarks below). In gall crabs, the parasitization of the pleon by species of
<taxonomicName id="4C1953BFFFB3AB56FC46FBBCE263FB7B" authorityName="Boyko &amp; van der Meij" authorityYear="2018" box="[956,1058,1076,1102]" class="Malacostraca" family="Bopyridae" genus="Spathione" kingdom="Animalia" order="Isopoda" pageId="9" pageNumber="156" phylum="Arthropoda" rank="genus">
<emphasis id="B96DF42EFFB3AB56FC46FBBCE263FB7B" box="[956,1058,1076,1102]" italics="true" pageId="9" pageNumber="156">Spathione</emphasis>
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may be a response to the host living in corals and making branchial parasitization more difficult. However, the soft carapace of pinnotherids and coral habitat of gall crabs do not preclude parasitization within the branchial chamber as four genera of keponine bopyrids (
<taxonomicName id="4C1953BFFFB3AB56FACAFB3DE580FBDB" authority="Shiino, 1936" authorityName="Shiino" authorityYear="1936" class="Malacostraca" family="Bopyridae" genus="Heterocepon" kingdom="Animalia" order="Isopoda" pageId="9" pageNumber="156" phylum="Arthropoda" rank="genus">
<emphasis id="B96DF42EFFB3AB56FACAFB3DE3EFFBFB" box="[1328,1454,1205,1230]" italics="true" pageId="9" pageNumber="156">Heterocepon</emphasis>
Shiino, 1936
</taxonomicName>
,
<taxonomicName id="4C1953BFFFB3AB56FC34FB5CE52BFA3B" authority="Nierstrasz &amp; Brender a Brandis, 1930" authorityName="Nierstrasz &amp; Brender a Brandis" authorityYear="1930" class="Malacostraca" family="Bopyridae" genus="Hypocepon" kingdom="Animalia" order="Isopoda" pageId="9" pageNumber="156" phylum="Arthropoda" rank="genus">
<emphasis id="B96DF42EFFB3AB56FC34FB5CE201FBD8" box="[974,1088,1236,1261]" italics="true" pageId="9" pageNumber="156">Hypocepon</emphasis>
<bibRefCitation id="EF8855CDFFB3AB56FBB3FB5CE52BFA3B" author="NIERSTRASZ H. F. &amp; BRENDER A BRANDIS G. A." pageId="9" pageNumber="156" pagination="12 - 17" refId="ref22593" refString="NIERSTRASZ H. F. &amp; BRENDER A BRANDIS G. A. 1930. - Resultats scientifiques du voyage aux Indes Orientales Neerlandaises de LL. AA. RR. le Prince et la Princesse Leopold de Belgique. Isopoda Epicaridea. Memoires du Musee royal d'Histoire naturelle de Belgique, Hors Serie 3 (1) (II): 12 - 17." type="journal article" year="1930">Nierstrasz &amp; Brender à Brandis, 1930</bibRefCitation>
</taxonomicName>
,
<taxonomicName id="4C1953BFFFB3AB56FC8EFB7BE2C6FA3B" authority="Perez, 1921" authorityName="Perez" authorityYear="1921" box="[884,1159,1267,1294]" class="Malacostraca" family="Bopyridae" genus="Onychocepon" kingdom="Animalia" order="Isopoda" pageId="9" pageNumber="156" phylum="Arthropoda" rank="genus">
<emphasis id="B96DF42EFFB3AB56FC8EFB7BE5BDFA38" box="[884,1020,1267,1293]" italics="true" pageId="9" pageNumber="156">Onychocepon</emphasis>
<bibRefCitation id="EF8855CDFFB3AB56FBFFFB7CE2C6FA3B" author="PEREZ C." box="[1029,1159,1268,1294]" pageId="9" pageNumber="156" pagination="59 - 61" refId="ref22961" refString="PEREZ C. 1921. - Sur un ceponien nouveau, Onychocepon harpax (n. g., n. sp.), parasite branchial d'un pinnothere. Comptes rendus hebdomadaires des seances de l'Academie des sciences 176 (2): 59 - 61." type="journal article" year="1921">Pérez, 1921</bibRefCitation>
</taxonomicName>
, and
<taxonomicName id="4C1953BFFFB3AB56FB3CFB7BE278FA1B" authority="Williams, Boyko &amp; Tri, 2023" authorityName="Williams, Boyko &amp; Tri" authorityYear="2023" class="Malacostraca" family="Bopyridae" genus="Rhizophoracepon" higherTaxonomySource="CoL" kingdom="Animalia" order="Isopoda" pageId="9" pageNumber="156" phylum="Arthropoda" rank="genus">
<emphasis id="B96DF42EFFB3AB56FB3CFB7BE334FA38" box="[1222,1397,1267,1293]" italics="true" pageId="9" pageNumber="156">Rhizophoracepon</emphasis>
<bibRefCitation id="EF8855CDFFB3AB56FA87FB7BE278FA1B" author="WILLIAMS J. D. &amp; BOYKO C. B. &amp; TRI N. V." pageId="9" pageNumber="156" pagination="531 - 541" refId="ref24339" refString="WILLIAMS J. D., BOYKO C. B. &amp; TRI N. V. 2023. - Description of a new genus and species of bopyrid (Isopoda: Epicaridea: Bopyridae) from the pinnotherid crab Plenotheres coarctatus (Burger, 1895) associated with mangrove clams from Vietnam. Raffles Bulletin of Zoology 71: 531 - 541. https: // doi. org / 10.26107 / RBZ- 2023 - 0040" type="journal article" year="2023">Williams, Boyko &amp;Tri, 2023</bibRefCitation>
</taxonomicName>
) are composed of entirely branchial bopyrids of pinnotherids, while a fourth genus (
<taxonomicName id="4C1953BFFFB3AB56FAD9FABBE590FA58" authority="Stebbing, 1910" authorityName="Stebbing" authorityYear="1910" class="Malacostraca" family="Bopyridae" genus="Dactylokepon" kingdom="Animalia" order="Isopoda" pageId="9" pageNumber="156" phylum="Arthropoda" rank="genus">
<emphasis id="B96DF42EFFB3AB56FAD9FABBE3EFFA78" box="[1315,1454,1331,1357]" italics="true" pageId="9" pageNumber="156">Dactylokepon</emphasis>
<bibRefCitation id="EF8855CDFFB3AB56FCD7FADBE590FA58" author="STEBBING T. R. R." box="[813,977,1363,1389]" pageId="9" pageNumber="156" pagination="83 - 122" refId="ref24077" refString="STEBBING T. R. R. 1910. - No. VI. - Isopoda from the Indian Ocean and British East Africa. Transactions of the Linnean Society of London, 2 nd Ser., Zoology 14 (1): 83 - 122, pls. 5 - 11. https: // doi. org / 10.1111 / j. 1096 - 3642.1910. tb 00525. x" type="journal article" year="1910">Stebbing, 1910</bibRefCitation>
</taxonomicName>
) contains one additional species known from pinnotherid branchial chambers. Likewise, the sole species of
<taxonomicName id="4C1953BFFFB3AB56FCB1FA1BE22FFA98" authority="Bourdon, 1983" authorityName="Bourdon" authorityYear="1983" box="[843,1134,1427,1453]" class="Malacostraca" family="Bopyridae" genus="Carcinione" kingdom="Animalia" order="Isopoda" pageId="9" pageNumber="156" phylum="Arthropoda" rank="genus">
<emphasis id="B96DF42EFFB3AB56FCB1FA1BE5FCFA98" box="[843,957,1427,1453]" italics="true" pageId="9" pageNumber="156">Carcinione</emphasis>
<bibRefCitation id="EF8855CDFFB3AB56FC3FFA1BE22FFA98" author="BOURDON R." box="[965,1134,1427,1453]" pageId="9" pageNumber="156" pagination="845 - 869" refId="ref19737" refString="BOURDON R. 1983. - Expedition Rumphius II (1975). Crustaces parasites, commensaux, etc. (Th. Monod Ed.). VIII. Crustaces isopodes (3 e partie: epicarides Bopyridae). Bulletin du Museum national d'Histoire naturelle, 4 e ser. 5 (A) (3): 845 - 869. https: // www. biodiversitylibrary. org / partpdf / 285968" type="journal article" year="1983">Bourdon, 1983</bibRefCitation>
</taxonomicName>
is a branchial parasite of gall crabs. Clearly, the mode of infestation exhibited by species of
<taxonomicName id="4C1953BFFFB3AB56FCD1FA5BE5DFFAD8" authorityName="Perez" authorityYear="1920" box="[811,926,1491,1517]" class="Malacostraca" family="Bopyridae" genus="Rhopalione" kingdom="Animalia" order="Isopoda" pageId="9" pageNumber="156" phylum="Arthropoda" rank="genus">
<emphasis id="B96DF42EFFB3AB56FCD1FA5BE5DFFAD8" box="[811,926,1491,1517]" italics="true" pageId="9" pageNumber="156">Rhopalione</emphasis>
</taxonomicName>
and
<taxonomicName id="4C1953BFFFB3AB56FC2FFA5BE27AFAD8" authorityName="Boyko &amp; van der Meij" authorityYear="2018" box="[981,1083,1491,1517]" class="Malacostraca" family="Bopyridae" genus="Spathione" kingdom="Animalia" order="Isopoda" pageId="9" pageNumber="156" phylum="Arthropoda" rank="genus">
<emphasis id="B96DF42EFFB3AB56FC2FFA5BE27AFAD8" box="[981,1083,1491,1517]" italics="true" pageId="9" pageNumber="156">Spathione</emphasis>
</taxonomicName>
is unusual among bopyrids found on pinnotherids and gall crabs but further discussion here is restricted to species of
<taxonomicName id="4C1953BFFFB3AB56FBE1F99AE2CFF919" authorityName="Perez" authorityYear="1920" box="[1051,1166,1554,1580]" class="Malacostraca" family="Bopyridae" genus="Rhopalione" kingdom="Animalia" order="Isopoda" pageId="9" pageNumber="156" phylum="Arthropoda" rank="genus">
<emphasis id="B96DF42EFFB3AB56FBE1F99AE2CFF919" box="[1051,1166,1554,1580]" italics="true" pageId="9" pageNumber="156">Rhopalione</emphasis>
</taxonomicName>
(see
<bibRefCitation id="EF8855CDFFB3AB56FB38F99AE524F978" author="BOYKO C. B. &amp; VAN DER MEIJ S. E. T." pageId="9" pageNumber="156" pagination="304 - 329" refId="ref20017" refString="BOYKO C. B. &amp; VAN DER MEIJ S. E. T. 2018. - A trifecta of Swiftian symbioses: stony corals, gall crabs and their parasites (Scleractinia; Brachyura: Cryptochiridae; Isopoda: Epicaridea and Cirripedia: Rhizocephala). Zoological Journal of the Linnean Society 184: 304 - 329. https: // doi. org / 10.1093 / zoolinnean / zlx 115" type="journal article" year="2018">Boyko &amp; van der Meij 2018</bibRefCitation>
for a fuller discussion on
<taxonomicName id="4C1953BFFFB3AB56FB86F9BAE2A3F979" authorityName="Boyko &amp; van der Meij" authorityYear="2018" box="[1148,1250,1586,1612]" class="Malacostraca" family="Bopyridae" genus="Spathione" kingdom="Animalia" order="Isopoda" pageId="9" pageNumber="156" phylum="Arthropoda" rank="genus">
<emphasis id="B96DF42EFFB3AB56FB86F9BAE2A3F979" box="[1148,1250,1586,1612]" italics="true" pageId="9" pageNumber="156">Spathione</emphasis>
</taxonomicName>
).
</paragraph>
<paragraph id="8BA6283CFFB3AB54FCB9F9DBE7A2FC25" blockId="9.[811,1457,375,2028]" lastBlockId="11.[130,777,215,1741]" lastPageId="11" lastPageNumber="158" pageId="9" pageNumber="156">
In other brachyuran hosts, the female parasites obtain food by piercing the gill lamellae and rarely leave evidence of their feeding. In contrast, females of species of
<taxonomicName id="4C1953BFFFB3AB56FB13F91AE31AF999" authorityName="Perez" authorityYear="1920" box="[1257,1371,1682,1708]" class="Malacostraca" family="Bopyridae" genus="Rhopalione" kingdom="Animalia" order="Isopoda" pageId="9" pageNumber="156" phylum="Arthropoda" rank="genus">
<emphasis id="B96DF42EFFB3AB56FB13F91AE31AF999" box="[1257,1371,1682,1708]" italics="true" pageId="9" pageNumber="156">Rhopalione</emphasis>
</taxonomicName>
are oriented with the dorsal surface appressed to the sternites of the host (
<bibRefCitation id="EF8855CDFFB3AB56FC9CF95AE590F9D8" author="PAGE R. D. M." box="[870,977,1746,1773]" pageId="9" pageNumber="156" pagination="185 - 212" refId="ref22839" refString="PAGE R. D. M. 1985. - Review of the New Zealand Bopyridae (Crustacea: Isopoda: Epicaridea). New Zealand Journal of Zoology 12: 185 - 212. https: // doi. org / 10.1080 / 03014223.1985.10428279" type="journal article" year="1985">Page 1985</bibRefCitation>
) and pierce the weakly calcified ventral surface of the pleon, leaving noticeable holes in the exoskeleton (see Remarks under
<taxonomicName id="4C1953BFFFB3AB56FC2CF89AE23AF81E" authority="Boyko &amp; Williams, 2024" authorityName="Boyko &amp; Williams" authorityYear="2024" box="[982,1147,1810,1836]" class="Malacostraca" family="Bopyridae" genus="Rhopalione" kingdom="Animalia" order="Isopoda" pageId="9" pageNumber="156" phylum="Arthropoda" rank="species" species="rusa" status="sp. nov.">
<emphasis id="B96DF42EFFB3AB56FC2CF89AE23AF81E" box="[982,1147,1810,1836]" italics="true" pageId="9" pageNumber="156">Rhopalione rusa</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A25E4955FFB3AB56FB78F89AE2FEF819" box="[1154,1215,1810,1836]" pageId="9" pageNumber="156" rank="species">n. sp.</taxonomicNameLabel>
). Females of species of
<taxonomicName id="4C1953BFFFB3AB56FCD7F8B9E5DEF87E" authorityName="Perez" authorityYear="1920" box="[813,927,1841,1867]" class="Malacostraca" family="Bopyridae" genus="Rhopalione" kingdom="Animalia" order="Isopoda" pageId="9" pageNumber="156" phylum="Arthropoda" rank="genus">
<emphasis id="B96DF42EFFB3AB56FCD7F8B9E5DEF87E" box="[813,927,1841,1867]" italics="true" pageId="9" pageNumber="156">Rhopalione</emphasis>
</taxonomicName>
are far more dependent on the secondary sexual modification of the host than other pseudionine species. Ordinarily, a female bopyrid occurring in a brachyuran branchial chamber is relatively unaffected by the sex of the host, as the host branchial chamber is not sexually dimorphic. However, in species of
<taxonomicName id="4C1953BFFFB3AB56FC4BF859E262F8DE" authorityName="Perez" authorityYear="1920" box="[945,1059,2001,2027]" class="Malacostraca" family="Bopyridae" genus="Rhopalione" kingdom="Animalia" order="Isopoda" pageId="9" pageNumber="156" phylum="Arthropoda" rank="genus">
<emphasis id="B96DF42EFFB3AB56FC4BF859E262F8DE" box="[945,1059,2001,2027]" italics="true" pageId="9" pageNumber="156">Rhopalione</emphasis>
</taxonomicName>
the parasite is found either on female crab hosts, which have a much broader pleon, or on male crab hosts that are usually feminized by the parasite, becoming modified with broader pleons (see
<bibRefCitation id="EF8855CDFFB1AB54FE3FFE90E477FE07" author="PAGE R. D. M." box="[453,566,279,306]" pageId="11" pageNumber="158" pagination="185 - 212" refId="ref22839" refString="PAGE R. D. M. 1985. - Review of the New Zealand Bopyridae (Crustacea: Isopoda: Epicaridea). New Zealand Journal of Zoology 12: 185 - 212. https: // doi. org / 10.1080 / 03014223.1985.10428279" type="journal article" year="1985">Page 1985</bibRefCitation>
). How the parasite accomplishes this feminization of the male host, which also has female-type pleopods accompanied by male gonopods (
<bibRefCitation id="EF8855CDFFB1AB54FD2EFEDFE6FDFEA4" author="PAGE R. D. M." pageId="11" pageNumber="158" pagination="185 - 212" refId="ref22839" refString="PAGE R. D. M. 1985. - Review of the New Zealand Bopyridae (Crustacea: Isopoda: Epicaridea). New Zealand Journal of Zoology 12: 185 - 212. https: // doi. org / 10.1080 / 03014223.1985.10428279" type="journal article" year="1985">Page 1985</bibRefCitation>
), is unknown but is likely hormonal in origin. Changes in sexual characters of male crab hosts has been documented in some bopyrids (e.g.
<bibRefCitation id="EF8855CDFFB1AB54FE8AFE3EE410FEE4" author="HIRAIWA Y. K. &amp; SATO M." box="[368,593,438,465]" pageId="11" pageNumber="158" pagination="105 - 124" refId="ref20766" refString="HIRAIWA Y. K. &amp; SATO M. 1939. - On the effect of parasitic Isopoda on a prawn, Penaeopsis akayebi Rathbun, with a consideration of the effect of parasitization on the higher Crustacea in general. Journal of Science of the Hiroshima University, Series B, Division 1 (Zoology) 7 (1 - 6): 105 - 124." type="journal article" year="1939">Hiraiwa &amp; Sato 1939</bibRefCitation>
;
<bibRefCitation id="EF8855CDFFB1AB54FDA4FE3EE541FEE4" author="REVERBERI G." box="[606,768,438,465]" pageId="11" pageNumber="158" pagination="56 - 70" refId="ref23304" refString="REVERBERI G. 1942. - Dati sulla trasformazione del sesso nei crostacei per opera del parassitismo da epicaridei. Pubblicazioni della Stazione Zoologica di Napoli 19: 56 - 70." type="journal article" year="1942">Reverberi 1942</bibRefCitation>
;
<bibRefCitation id="EF8855CDFFB1AB54FF7EFE5EE75FFEC4" author="REINHARD E. G." box="[132,286,470,497]" pageId="11" pageNumber="158" pagination="79 - 107" refId="ref23262" refString="REINHARD E. G. 1956. - Parasitic castration of Crustacea. Experimental Parasitology 5 (1): 79 - 107. https: // doi. org / 10.1016 / 0014 - 4894 (56) 90007 - 8" type="journal article" year="1956">Reinhard 1956</bibRefCitation>
;
<bibRefCitation id="EF8855CDFFB1AB54FED0FE5EE780FEC5" author="BOURDON R." box="[298,449,470,496]" pageId="11" pageNumber="158" pagination="77 - 424" refId="ref19586" refString="BOURDON R. 1968. - Les Bopyridae des mers europeennes. Museum national d'Histoire naturelle, Paris (Memoires du Museum national d'Histoire naturelle, Ser. A - Zoologie (1950 - 1992); 50 (2): 77 - 424. https: // www. biodiversitylibrary. org / part / 280651" type="journal article" year="1968">Bourdon 1968</bibRefCitation>
; Romero-Rodríguez &amp; RománContreras 2011) but is usually limited to primary reproductive structures (gonopores, gonopods, pleopods) and with a much less pronounced effect on external morphology and secondary sexual characters. It is also highly variable, as all hosts are not equally affected by their parasites in terms of modification of sexual characters. A side-effect of the abdominal mode of life is that no specimens of any species of
<taxonomicName id="4C1953BFFFB1AB54FDB9FD3DE4F4FDFA" authorityName="Perez" authorityYear="1920" box="[579,693,693,719]" class="Malacostraca" family="Bopyridae" genus="Rhopalione" kingdom="Animalia" order="Isopoda" pageId="11" pageNumber="158" phylum="Arthropoda" rank="genus">
<emphasis id="B96DF42EFFB1AB54FDB9FD3DE4F4FDFA" box="[579,693,693,719]" italics="true" pageId="11" pageNumber="158">Rhopalione</emphasis>
</taxonomicName>
exhibit marked torsion of the body (dextral or sinistral), as commonly seen in many other bopyrid taxa.
</paragraph>
<caption id="DF6678B4FFB0AB55FF7EF81BE5D0F8DC" ID-DOI="http://doi.org/10.5281/zenodo.10863459" ID-Zenodo-Dep="10863459" httpUri="https://zenodo.org/record/10863459/files/figure.png" pageId="10" pageNumber="157" startId="10.[132,143,1939,1957]" targetBox="[132,1455,93,1823]" targetPageId="10" targetType="figure">
<paragraph id="8BA6283CFFB0AB55FF7EF81BE5D0F8DC" blockId="10.[132,1457,1939,2025]" pageId="10" pageNumber="157">
<smallCapsWord id="8D40BEE0FFB0AB55FF7EF81BE6DCF891" baselines="1952,1953" box="[132,157,1939,1957]" lowerCaseFontSize="5" mainFontSize="7" normCase="title" normString="Fig" pageId="10" pageNumber="157">FIG</smallCapsWord>
. 3. —
<taxonomicName id="4C1953BFFFB0AB55FF2EF81BE7B1F890" authority="Perez, 1921" authorityName="Perez" authorityYear="1921" box="[212,496,1939,1957]" class="Malacostraca" family="Bopyridae" genus="Onychocepon" kingdom="Animalia" order="Isopoda" pageId="10" pageNumber="157" phylum="Arthropoda" rank="species" species="harpax">
<emphasis id="B96DF42EFFB0AB55FF2EF81BE7C7F890" box="[212,390,1939,1957]" italics="true" pageId="10" pageNumber="157">Onychocepon harpax</emphasis>
<bibRefCitation id="EF8855CDFFB0AB55FE71F81BE7B1F890" author="PEREZ C." box="[395,496,1939,1957]" pageId="10" pageNumber="157" pagination="59 - 61" refId="ref22961" refString="PEREZ C. 1921. - Sur un ceponien nouveau, Onychocepon harpax (n. g., n. sp.), parasite branchial d'un pinnothere. Comptes rendus hebdomadaires des seances de l'Academie des sciences 176 (2): 59 - 61." type="journal article" year="1921">Pérez, 1921</bibRefCitation>
</taxonomicName>
, ZRC 2023.0258 (
<emphasis id="B96DF42EFFB0AB55FD7FF81BE4E7F890" bold="true" box="[645,678,1939,1957]" pageId="10" pageNumber="157">A -K</emphasis>
):
<emphasis id="B96DF42EFFB0AB55FD4CF81BE482F890" bold="true" box="[694,707,1939,1957]" pageId="10" pageNumber="157">A</emphasis>
, female, dorsal view;
<emphasis id="B96DF42EFFB0AB55FC82F81BE5C4F890" bold="true" box="[888,901,1939,1957]" pageId="10" pageNumber="157">B</emphasis>
, male, dorsal view;
<emphasis id="B96DF42EFFB0AB55FBD0F81BE279F890" bold="true" box="[1066,1080,1939,1957]" pageId="10" pageNumber="157">C</emphasis>
, male, ventral view (arrows indicate attached ciliates);
<emphasis id="B96DF42EFFB0AB55FF34F822E69DF889" bold="true" box="[206,220,1962,1980]" pageId="10" pageNumber="157">D</emphasis>
, left antennule of female;
<emphasis id="B96DF42EFFB0AB55FE43F822E784F889" bold="true" box="[441,453,1962,1980]" pageId="10" pageNumber="157">E</emphasis>
, left antenna of female;
<emphasis id="B96DF42EFFB0AB55FD69F822E4DFF889" bold="true" box="[659,670,1962,1980]" pageId="10" pageNumber="157">F</emphasis>
, left maxilliped and barbular lobes of female;
<emphasis id="B96DF42EFFB0AB55FBD8F822E271F889" bold="true" box="[1058,1072,1962,1980]" pageId="10" pageNumber="157">G</emphasis>
, left oostegite 1 of female, outer view;
<emphasis id="B96DF42EFFB0AB55FA86F822E3CBF889" bold="true" box="[1404,1418,1962,1980]" pageId="10" pageNumber="157">H</emphasis>
, left oostegite 1 of female, inner view, asterisk shows lobe drawn in close-up;
<emphasis id="B96DF42EFFB0AB55FD26F849E4A3F8E6" bold="true" box="[732,738,1985,2003]" pageId="10" pageNumber="157">I</emphasis>
, close-up of lobe from inner ridge of oostegite 1 of female;
<emphasis id="B96DF42EFFB0AB55FB32F849E293F8E6" bold="true" box="[1224,1234,1985,2003]" pageId="10" pageNumber="157">J</emphasis>
, left pereopod 1 of female;
<emphasis id="B96DF42EFFB0AB55FF7EF85FE6D0F8DC" bold="true" box="[132,145,2007,2025]" pageId="10" pageNumber="157">K</emphasis>
, left pereopod 7 of female. Scale bars: A, 2 mm; B, C, 1 mm; D, E, I-K, 50 µm; F-H, 200 µm.
</paragraph>
</caption>
<paragraph id="8BA6283CFFB1AB54FF61FC9DE406F9F8" blockId="11.[130,777,215,1741]" pageId="11" pageNumber="158">
The relationships of
<taxonomicName id="4C1953BFFFB1AB54FE7FFC9DE7BCFC1A" authorityName="Perez" authorityYear="1920" box="[389,509,789,815]" class="Malacostraca" family="Bopyridae" genus="Rhopalione" kingdom="Animalia" order="Isopoda" pageId="11" pageNumber="158" phylum="Arthropoda" rank="genus">
<emphasis id="B96DF42EFFB1AB54FE7FFC9DE7BCFC1A" box="[389,509,789,815]" italics="true" pageId="11" pageNumber="158">Rhopalione</emphasis>
</taxonomicName>
and
<taxonomicName id="4C1953BFFFB1AB54FDC3FC9DE4E2FC1A" authorityName="Boyko &amp; van der Meij" authorityYear="2018" box="[569,675,789,815]" class="Malacostraca" family="Bopyridae" genus="Spathione" kingdom="Animalia" order="Isopoda" pageId="11" pageNumber="158" phylum="Arthropoda" rank="genus">
<emphasis id="B96DF42EFFB1AB54FDC3FC9DE4E2FC1A" box="[569,675,789,815]" italics="true" pageId="11" pageNumber="158">Spathione</emphasis>
</taxonomicName>
to other pseudionine bopyrids are difficult to discern.
<taxonomicName id="4C1953BFFFB1AB54FD93FCBDE49DFC7A" authorityName="Perez" authorityYear="1920" box="[617,732,821,847]" class="Malacostraca" family="Bopyridae" genus="Rhopalione" kingdom="Animalia" order="Isopoda" pageId="11" pageNumber="158" phylum="Arthropoda" rank="genus">
<emphasis id="B96DF42EFFB1AB54FD93FCBDE49DFC7A" box="[617,732,821,847]" italics="true" pageId="11" pageNumber="158">Rhopalione</emphasis>
</taxonomicName>
was once considered a part of Ioninae H. Milne Edwards, 1840
<emphasis id="B96DF42EFFB1AB54FF7FFCFEE6A5FCBA" box="[133,228,885,911]" italics="true" pageId="11" pageNumber="158">sensu lato</emphasis>
(now
<taxonomicName id="4C1953BFFFB1AB54FEDFFCFDE6FAFC9A" authority="Boyko, Moss, Williams &amp; Shields, 2013" authorityName="Boyko, Moss, Williams &amp; Shields" authorityYear="2013" class="Malacostraca" family="Bopyridae" kingdom="Animalia" order="Isopoda" pageId="11" pageNumber="151" phylum="Arthropoda" rank="subFamily" subFamily="Keponinae">
Keponinae
<bibRefCitation id="EF8855CDFFB1AB54FE64FCFDE6FAFC9A" author="BOYKO C. B. &amp; MOSS J. &amp; WILLIAMS J. D. &amp; SHIELDS J. D." pageId="11" pageNumber="158" pagination="495 - 506" refId="ref19961" refString="BOYKO C. B., MOSS J., WILLIAMS J. D. &amp; SHIELDS J. D. 2013. - A molecular phylogeny of Bopyroidea and Cryptoniscoidea (Crustacea: Isopoda). Systematics and Biodiversity 11: 495 - 506. https: // doi. org / 10.1080 / 14772000.2013.865679" type="journal article" year="2013">Boyko, Moss, Williams &amp; Shields, 2013</bibRefCitation>
</taxonomicName>
) but was moved to Pseudioninae Codreanu, 1967 by
<bibRefCitation id="EF8855CDFFB1AB54FF7EFC3DE782FCFA" author="BOYKO C. B. &amp; VAN DER MEIJ S. E. T." box="[132,451,949,975]" pageId="11" pageNumber="158" pagination="304 - 329" refId="ref20017" refString="BOYKO C. B. &amp; VAN DER MEIJ S. E. T. 2018. - A trifecta of Swiftian symbioses: stony corals, gall crabs and their parasites (Scleractinia; Brachyura: Cryptochiridae; Isopoda: Epicaridea and Cirripedia: Rhizocephala). Zoological Journal of the Linnean Society 184: 304 - 329. https: // doi. org / 10.1093 / zoolinnean / zlx 115" type="journal article" year="2018">Boyko &amp; van der Meij (2018)</bibRefCitation>
, as the pleonal lateral plates of the female are very well developed and proximally constricted, the dorsomedial regions of the pereomeres are raised into low bosses, and the maxilliped and oostegites are comparable to other genera in the subfamily.Within Pseudioninae, however, these two genera are difficult to place, as they bear no general resemblance to any other taxon. The males of
<taxonomicName id="4C1953BFFFB1AB54FD73FBFCE4BFFBBB" authorityName="Perez" authorityYear="1920" box="[649,766,1140,1166]" class="Malacostraca" family="Bopyridae" genus="Rhopalione" kingdom="Animalia" order="Isopoda" pageId="11" pageNumber="158" phylum="Arthropoda" rank="genus">
<emphasis id="B96DF42EFFB1AB54FD73FBFCE4BFFBBB" box="[649,766,1140,1166]" italics="true" pageId="11" pageNumber="158">Rhopalione</emphasis>
</taxonomicName>
, with their distinctive pleonal morphology and strong dorsoventral curvature, are also unlike other pseudionine species, and indeed other bopyrid taxa excepting
<taxonomicName id="4C1953BFFFB1AB54FDCEFB5CE4DBFBDB" authorityName="Boyko &amp; van der Meij" authorityYear="2018" box="[564,666,1236,1262]" class="Malacostraca" family="Bopyridae" genus="Spathione" kingdom="Animalia" order="Isopoda" pageId="11" pageNumber="158" phylum="Arthropoda" rank="genus">
<emphasis id="B96DF42EFFB1AB54FDCEFB5CE4DBFBDB" box="[564,666,1236,1262]" italics="true" pageId="11" pageNumber="158">Spathione</emphasis>
</taxonomicName>
.
<bibRefCitation id="EF8855CDFFB1AB54FD5CFB5CE68FFA3B" author="BOURDON R." pageId="11" pageNumber="158" pagination="845 - 869" refId="ref19737" refString="BOURDON R. 1983. - Expedition Rumphius II (1975). Crustaces parasites, commensaux, etc. (Th. Monod Ed.). VIII. Crustaces isopodes (3 e partie: epicarides Bopyridae). Bulletin du Museum national d'Histoire naturelle, 4 e ser. 5 (A) (3): 845 - 869. https: // www. biodiversitylibrary. org / partpdf / 285968" type="journal article" year="1983">Bourdon (1983)</bibRefCitation>
cited
<taxonomicName id="4C1953BFFFB1AB54FEEFFB7BE7C8FA38" authorityName="Perez" authorityYear="1920" box="[277,393,1267,1293]" class="Malacostraca" family="Bopyridae" genus="Rhopalione" kingdom="Animalia" order="Isopoda" pageId="11" pageNumber="158" phylum="Arthropoda" rank="genus">
<emphasis id="B96DF42EFFB1AB54FEEFFB7BE7C8FA38" box="[277,393,1267,1293]" italics="true" pageId="11" pageNumber="158">Rhopalione</emphasis>
</taxonomicName>
,
<taxonomicName id="4C1953BFFFB1AB54FE63FB7CE490FA3B" authority="Kossmann, 1881" authorityName="Kossmann" authorityYear="1881" box="[409,721,1268,1294]" class="Malacostraca" family="Bopyridae" genus="Gigantione" kingdom="Animalia" order="Isopoda" pageId="11" pageNumber="158" phylum="Arthropoda" rank="genus">
<emphasis id="B96DF42EFFB1AB54FE63FB7CE44CFA3B" box="[409,525,1268,1294]" italics="true" pageId="11" pageNumber="158">Gigantione</emphasis>
<bibRefCitation id="EF8855CDFFB1AB54FDEFFB7CE490FA3B" author="KOSSMANN R." box="[533,721,1268,1294]" pageId="11" pageNumber="158" pagination="652 - 680" refId="ref21115" refString="KOSSMANN R. 1881. - Studien uber bopyriden. Zeitschrift fur Wissenschaftliche Zoologie 35 (4): 652 - 680, pls. 32 - 35. https: // www. biodiversitylibrary. org / part / 186356" type="journal article" year="1881">Kossmann, 1881</bibRefCitation>
</taxonomicName>
, and
<taxonomicName id="4C1953BFFFB1AB54FF7CFA9CE7E4FA1B" authority="Bourdon, 1983" authorityName="Bourdon" authorityYear="1983" box="[134,421,1299,1326]" class="Malacostraca" family="Bopyridae" genus="Carcinione" kingdom="Animalia" order="Isopoda" pageId="11" pageNumber="158" phylum="Arthropoda" rank="genus">
<emphasis id="B96DF42EFFB1AB54FF7CFA9CE6B9FA1B" box="[134,248,1300,1326]" italics="true" pageId="11" pageNumber="158">Carcinione</emphasis>
<bibRefCitation id="EF8855CDFFB1AB54FF07FA9BE7E4FA1B" author="BOURDON R." box="[253,421,1299,1326]" pageId="11" pageNumber="158" pagination="845 - 869" refId="ref19737" refString="BOURDON R. 1983. - Expedition Rumphius II (1975). Crustaces parasites, commensaux, etc. (Th. Monod Ed.). VIII. Crustaces isopodes (3 e partie: epicarides Bopyridae). Bulletin du Museum national d'Histoire naturelle, 4 e ser. 5 (A) (3): 845 - 869. https: // www. biodiversitylibrary. org / partpdf / 285968" type="journal article" year="1983">Bourdon, 1983</bibRefCitation>
</taxonomicName>
, as the only three genera infesting brachyurans that did not appear to belong to Ioninae, but he did not formally assign any of them to a subfamily. However,
<bibRefCitation id="EF8855CDFFB1AB54FF7EFAFBE702FAB8" author="BOURDON R." box="[132,323,1395,1421]" pageId="11" pageNumber="158" pagination="845 - 869" refId="ref19737" refString="BOURDON R. 1983. - Expedition Rumphius II (1975). Crustaces parasites, commensaux, etc. (Th. Monod Ed.). VIII. Crustaces isopodes (3 e partie: epicarides Bopyridae). Bulletin du Museum national d'Histoire naturelle, 4 e ser. 5 (A) (3): 845 - 869. https: // www. biodiversitylibrary. org / partpdf / 285968" type="journal article" year="1983">Bourdons (1983)</bibRefCitation>
consideration of the status of
<taxonomicName id="4C1953BFFFB1AB54FD69FAFBE547FAB8" authorityName="Perez" authorityYear="1920" box="[659,774,1395,1421]" class="Malacostraca" family="Bopyridae" genus="Rhopalione" kingdom="Animalia" order="Isopoda" pageId="11" pageNumber="158" phylum="Arthropoda" rank="genus">
<emphasis id="B96DF42EFFB1AB54FD69FAFBE547FAB8" box="[659,774,1395,1421]" italics="true" pageId="11" pageNumber="158">Rhopalione</emphasis>
</taxonomicName>
was based solely on information in
<bibRefCitation id="EF8855CDFFB1AB54FE0BFA1BE4C0FA98" author="PEREZ C." box="[497,641,1427,1453]" pageId="11" pageNumber="158" pagination="1615 - 1617" refId="ref22884" refString="PEREZ C. 1920 a. - Sur un type nouveau d'epicarides, Rhopalione uromyzon n. g. n. sp. parasite sous-abdominal d'un Pinnothere Comptes rendus hebdomadaires des seances de l'Academie des sciences 170: 1615 - 1617. https: // www. biodiversitylibrary. org / partpdf / 92296" type="journal article" year="1920">Pérez (1920a)</bibRefCitation>
. Subsequent work by
<bibRefCitation id="EF8855CDFFB1AB54FF1DFA3BE72AFAFB" author="PAGE R. D. M." box="[231,363,1459,1486]" pageId="11" pageNumber="158" pagination="185 - 212" refId="ref22839" refString="PAGE R. D. M. 1985. - Review of the New Zealand Bopyridae (Crustacea: Isopoda: Epicaridea). New Zealand Journal of Zoology 12: 185 - 212. https: // doi. org / 10.1080 / 03014223.1985.10428279" type="journal article" year="1985">Page (1985)</bibRefCitation>
and
<bibRefCitation id="EF8855CDFFB1AB54FE5FFA3BE422FAF8" author="MARKHAM J. C." box="[421,611,1459,1485]" pageId="11" pageNumber="158" pagination="555 - 568" refId="ref21666" refString="MARKHAM J. C. 1990. - Further notes on the Isopoda Bopyridae of Hong Kong, in MORTON B. (ed.), Proceedings of the Second International Marine Biological Workshop: The Marine Flora and Fauna of Hong Kong and Southern China, Hong Kong, 1986 2: 555 - 568." type="journal article" year="1990">Markham (1990)</bibRefCitation>
suggested that
<taxonomicName id="4C1953BFFFB1AB54FF7CFA5BE6BDFAD8" authorityName="Perez" authorityYear="1920" box="[134,252,1491,1517]" class="Malacostraca" family="Bopyridae" genus="Rhopalione" kingdom="Animalia" order="Isopoda" pageId="11" pageNumber="158" phylum="Arthropoda" rank="genus">
<emphasis id="B96DF42EFFB1AB54FF7CFA5BE6BDFAD8" box="[134,252,1491,1517]" italics="true" pageId="11" pageNumber="158">Rhopalione</emphasis>
</taxonomicName>
was a member of Ioninae (of uncertain placement) but this was refuted by
<bibRefCitation id="EF8855CDFFB1AB54FE3CFA7BE546F938" author="BOYKO C. B. &amp; VAN DER MEIJ S. E. T." box="[454,775,1523,1549]" pageId="11" pageNumber="158" pagination="304 - 329" refId="ref20017" refString="BOYKO C. B. &amp; VAN DER MEIJ S. E. T. 2018. - A trifecta of Swiftian symbioses: stony corals, gall crabs and their parasites (Scleractinia; Brachyura: Cryptochiridae; Isopoda: Epicaridea and Cirripedia: Rhizocephala). Zoological Journal of the Linnean Society 184: 304 - 329. https: // doi. org / 10.1093 / zoolinnean / zlx 115" type="journal article" year="2018">Boyko &amp; van der Meij (2018)</bibRefCitation>
who demonstrated that the genus belongs to Pseudioninae.
<taxonomicName id="4C1953BFFFB1AB54FF7FF9BBE6B9F978" authorityName="Kossmann" authorityYear="1881" box="[133,248,1587,1613]" class="Malacostraca" family="Bopyridae" genus="Gigantione" kingdom="Animalia" order="Isopoda" pageId="11" pageNumber="158" phylum="Arthropoda" rank="genus">
<emphasis id="B96DF42EFFB1AB54FF7FF9BBE6B9F978" box="[133,248,1587,1613]" italics="true" pageId="11" pageNumber="158">Gigantione</emphasis>
</taxonomicName>
may be paraphyletic (Boyko, unpublished) but its position, as defined by the
<typeStatus id="54A2969EFFB1AB54FE65F9DBE78CF958" box="[415,461,1619,1645]" pageId="11" pageNumber="158">type</typeStatus>
species
<taxonomicName id="4C1953BFFFB1AB54FDDBF9DBE6FDF9B9" authority="Kossmann, 1881" authorityName="Kossmann" authorityYear="1881" class="Malacostraca" family="Bopyridae" genus="Gigantione" kingdom="Animalia" order="Isopoda" pageId="11" pageNumber="158" phylum="Arthropoda" rank="species" species="moebii">
<emphasis id="B96DF42EFFB1AB54FDDBF9DBE474F958" box="[545,565,1619,1645]" italics="true" pageId="11" pageNumber="158">G</emphasis>
.
<emphasis id="B96DF42EFFB1AB54FDB9F9DAE4C8F959" box="[579,649,1618,1644]" italics="true" pageId="11" pageNumber="158">moebii</emphasis>
<bibRefCitation id="EF8855CDFFB1AB54FD75F9DBE6FDF9B9" author="KOSSMANN R." pageId="11" pageNumber="158" pagination="652 - 680" refId="ref21115" refString="KOSSMANN R. 1881. - Studien uber bopyriden. Zeitschrift fur Wissenschaftliche Zoologie 35 (4): 652 - 680, pls. 32 - 35. https: // www. biodiversitylibrary. org / part / 186356" type="journal article" year="1881">Kossmann, 1881</bibRefCitation>
</taxonomicName>
(see also
<bibRefCitation id="EF8855CDFFB1AB54FED3F9FAE78CF9B9" author="BOURDON R." box="[297,461,1650,1676]" pageId="11" pageNumber="158" pagination="947 - 952" refId="ref19644" refString="BOURDON R. 1969. - Xanthias punctatus (H. Milne Edwards), nouvel hote pour Gigantione moebii Kossmann. Revue Suisse de Zoologie 76 (48): 947 - 952. https: // doi. org / 10.5962 / bhl. part. 97733" type="journal article" year="1969">Bourdon 1969</bibRefCitation>
), suggests placement within Pseudioninae as well.
<taxonomicName id="4C1953BFFFB1AB54FE8DF91BE7A8F998" authorityName="Bourdon" authorityYear="1983" box="[375,489,1683,1709]" class="Malacostraca" family="Bopyridae" genus="Carcinione" kingdom="Animalia" order="Isopoda" pageId="11" pageNumber="158" phylum="Arthropoda" rank="genus">
<emphasis id="B96DF42EFFB1AB54FE8DF91BE7A8F998" box="[375,489,1683,1709]" italics="true" pageId="11" pageNumber="158">Carcinione</emphasis>
</taxonomicName>
was also placed in Pseudioninae by
<bibRefCitation id="EF8855CDFFB1AB54FF06F93AE400F9F9" author="BOYKO C. B. &amp; VAN DER MEIJ S. E. T." box="[252,577,1714,1741]" pageId="11" pageNumber="158" pagination="304 - 329" refId="ref20017" refString="BOYKO C. B. &amp; VAN DER MEIJ S. E. T. 2018. - A trifecta of Swiftian symbioses: stony corals, gall crabs and their parasites (Scleractinia; Brachyura: Cryptochiridae; Isopoda: Epicaridea and Cirripedia: Rhizocephala). Zoological Journal of the Linnean Society 184: 304 - 329. https: // doi. org / 10.1093 / zoolinnean / zlx 115" type="journal article" year="2018">Boyko &amp; van der Meij (2018)</bibRefCitation>
.
</paragraph>
</subSubSection>
</treatment>
</document>