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<mods:title>Hydromedusae observed during night dives in the Gulf Stream</mods:title>
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<mods:namePart>Schuchert, Peter</mods:namePart>
<mods:affiliation>Muséum d’histoire naturelle, C. P. 6434, CH- 1211 Genève 6, Switzerland</mods:affiliation>
<mods:nameIdentifier type="email">peter.schuchert@ville-ge.ch</mods:nameIdentifier>
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<mods:namePart>Collins, Richard</mods:namePart>
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<taxonomicName ID-CoL="6ZHV4" ID-ENA="2805962" authority="Haeckel, 1879" authorityName="Haeckel" authorityYear="1879" box="[231,676,1241,1266]" class="Hydrozoa" family="Hebellidae" genus="Staurodiscus" kingdom="Animalia" order="Leptothecata" pageId="51" pageNumber="287" phylum="Cnidaria" rank="species" species="tetrastaurus">
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<emphasis bold="true" box="[231,511,1241,1265]" italics="true" pageId="51" pageNumber="287">Staurodiscus tetrastaurus</emphasis>
<bibRefCitation author="Haeckel" box="[518,676,1242,1266]" firstAuthor="Haeckel" pageId="51" pageNumber="287" pagination="1 - 360" refId="ref68751" refString="Haeckel E. 1879. Das System der Medusen. Erster Teil einer Monographie der Medusen. Denkschriften der Medicinisch- Naturwissenschaftlichen Gesellschaft zu Jena 1: I-XX, 1 - 360, 20 pls." type="book chapter" year="1879">Haeckel, 1879</bibRefCitation>
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<paragraph blockId="51.[231,676,1241,1298]" box="[390,517,1274,1298]" pageId="51" pageNumber="287">
<figureCitation box="[390,469,1274,1298]" captionStart="Fig" captionStartId="52.[151,186,1998,2020]" captionTargetBox="[151,1436,648,1932]" captionTargetId="figure-165@52.[151,1436,648,1932]" captionTargetPageId="52" captionText="Fig. 32. Staurodiscus tetrastaurus. (A-C) Young animal observed 11-DEC-2017. (D-E) BFLA3831, size 7 mm. Some amphipods are associated with the animal. (F) BFLA4440, size 10 mm, bell margin seen from adaxial side, ocelli are barely visible. (G) BFLA4440, oral view, note variation of position of lateral diverticula." figureDoi="http://doi.org/10.5281/zenodo.5640032" httpUri="https://zenodo.org/record/5640032/files/figure.png" pageId="51" pageNumber="287">Figs 32</figureCitation>
A-D
</paragraph>
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<treatmentCitation author="Haeckel E." box="[151,596,1319,1341]" page="145" pageId="51" pageNumber="287" year="1879">
<taxonomicName ID-CoL="6ZHV4" authority="Haeckel, 1879: 145" authorityName="Haeckel" authorityPageNumber="145" authorityYear="1879" box="[151,596,1319,1341]" class="Hydrozoa" family="Hebellidae" genus="Staurodiscus" kingdom="Animalia" order="Leptothecata" pageId="51" pageNumber="287" phylum="Cnidaria" rank="species" species="tetrastaurus">
<emphasis box="[151,397,1319,1341]" italics="true" pageId="51" pageNumber="287">Staurodiscus tetrastaurus</emphasis>
<bibRefCitation author="Haeckel" box="[404,596,1319,1341]" firstAuthor="Haeckel" page="145" pageId="51" pageNumber="287" pagination="1 - 360" refId="ref68751" refString="Haeckel E. 1879. Das System der Medusen. Erster Teil einer Monographie der Medusen. Denkschriften der Medicinisch- Naturwissenschaftlichen Gesellschaft zu Jena 1: I-XX, 1 - 360, 20 pls." type="book chapter" year="1879">Haeckel, 1879: 145</bibRefCitation>
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, pl. 9 figs 1-3. – 
<treatmentCitation author="Mayer A. G." box="[227,403,1349,1370]" page="214" pageId="51" pageNumber="287" year="1910">
<bibRefCitation author="Mayer" box="[227,403,1349,1370]" firstAuthor="Mayer" page="214" pageId="51" pageNumber="287" pagination="1 - 76" refId="ref71665" refString="Mayer A. G. 1910. Medusae of the world. Hydromedusae, Vols. I &amp; II. Scyphomedusae, Vol III. Carnegie Institution, Washington, pp. 735, plates 1 - 76." type="journal article" year="1910">Mayer, 1910: 214</bibRefCitation>
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, pl. 22 figs 7-8, pl. 25 fig. 5, pl. 26 figs 10-11. – 
<treatmentCitation author="Uchida T." box="[357,553,1378,1400]" page="167" pageId="51" pageNumber="287" year="1927">
<bibRefCitation author="Uchida" box="[357,553,1378,1400]" firstAuthor="Uchida" page="167" pageId="51" pageNumber="287" pagination="165 - 168" refId="ref75634" refString="Uchida T. 1927 b. Description of a new Leptomedusa, Staurodiscoides gotoi. Japanese Journal of Zoology 1 (5): 165 - 168." type="journal article" year="1927" yearSuffix="b">Uchida, 1927b: 167</bibRefCitation>
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. – 
<treatmentCitation author="Menon M. G. K." box="[582,752,1378,1399]" page="14" pageId="51" pageNumber="287" year="1932">
<bibRefCitation author="Menon" box="[582,752,1378,1399]" firstAuthor="Menon" page="14" pageId="51" pageNumber="287" pagination="1 - 32" refId="ref71864" refString="Menon M. G. K. 1932. The Hydromedusae of Madras. Bulletin of the Madras Government Museum, new series, Natural History Section 3 (2): 1 - 32, pls 1 - 3." type="journal article" year="1932">Menon, 1932: 14</bibRefCitation>
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, pl. 2 figs 14, 19. – 
<treatmentCitation author="Nair K. K." box="[401,540,1407,1429]" page="60" pageId="51" pageNumber="287" year="1951">
<bibRefCitation author="Nair" box="[401,540,1407,1429]" firstAuthor="Nair" page="60" pageId="51" pageNumber="287" pagination="47 - 75" refId="ref72659" refString="Nair K. K. 1951. Medusae of the Trivandrum Coast. Part I. Systematics. Bulletin of the Research Institute of the University of Travancore 20 (1): 47 - 75, pl. 1." type="journal article" year="1951">Nair, 1951: 60</bibRefCitation>
</treatmentCitation>
. – 
<treatmentCitation author="Kramp P. L." box="[566,752,1407,1429]" page="140" pageId="51" pageNumber="287" year="1959">
<bibRefCitation author="Kramp" box="[566,752,1407,1429]" firstAuthor="Kramp" page="140" pageId="51" pageNumber="287" pagination="1 - 283" refId="ref69812" refString="Kramp P. L. 1959 a. The Hydromedusae of the Atlantic Ocean and adjacent waters. Dana Report 46: 1 - 283." type="book chapter" year="1959" yearSuffix="a">Kramp, 1959a: 140</bibRefCitation>
</treatmentCitation>
, fig. 165. – 
<treatmentCitation author="Kramp P. L." box="[336,519,1436,1458]" page="148" pageId="51" pageNumber="287" year="1961">
<bibRefCitation author="Kramp" box="[336,519,1436,1458]" firstAuthor="Kramp" page="148" pageId="51" pageNumber="287" pagination="1 - 469" refId="ref69868" refString="Kramp P. L. 1961. Synopsis of the medusae of the world. Journal of the Marine Biological Association of the U. K. 40: 1 - 469." type="journal article" year="1961">Kramp, 1961: 148</bibRefCitation>
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. – 
<treatmentCitation author="Kramp P. L." box="[550,733,1436,1458]" page="321" pageId="51" pageNumber="287" year="1962">
<bibRefCitation author="Kramp" box="[550,733,1436,1458]" firstAuthor="Kramp" page="321" pageId="51" pageNumber="287" pagination="305 - 366" refId="ref69901" refString="Kramp P. L. 1962. Medusae of Vietnam. Videnskabelige Meddelelser fra Dansk naturhistorisk Forening i KObenhavn 124: 305 - 366." type="journal article" year="1962">Kramp, 1962: 321</bibRefCitation>
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. – 
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<bibRefCitation author="Kramp" box="[227,391,1465,1487]" firstAuthor="Kramp" page="70" pageId="51" pageNumber="287" pagination="1 - 200" refId="ref69950" refString="Kramp P. L. 1968. The hydromedusae of the Pacific and Indian oceans. Sections II and III. Dana Report 72: 1 - 200." type="book chapter" year="1968">Kramp, 1968: 70</bibRefCitation>
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, fig. 182. – 
<treatmentCitation author="Migotto A. E. &amp; De Andrade L. P." box="[502,753,1465,1487]" pageId="51" pageNumber="287" year="2000">
<bibRefCitation author="Migotto &amp; Andrade" box="[502,753,1465,1487]" firstAuthor="Migotto" pageId="51" pageNumber="287" pagination="1871 - 1888" refId="ref72189" refString="Migotto A. E., De Andrade L. P. 2000. The life cycle of Hebella furax (Cnidaria: Hydrozoa): a link between a lafoeid hydroid and a laodiceid medusa. Journal of natural History 34 (10): 1871 - 1888." type="journal article" year="2000">Migotto &amp; Andrade, 2000</bibRefCitation>
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: table 6.
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</paragraph>
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<treatmentCitationGroup box="[151,692,1523,1545]" pageId="51" pageNumber="287">
<treatmentCitation author="Haeckel E." box="[151,594,1523,1545]" page="146" pageId="51" pageNumber="287" year="1879">
<taxonomicName ID-CoL="4ZM4K" authority="Haeckel, 1879: 146" authorityName="Haeckel" authorityPageNumber="146" authorityYear="1879" box="[151,594,1523,1545]" class="Hydrozoa" family="Hebellidae" genus="Staurodiscus" kingdom="Animalia" order="Leptothecata" pageId="51" pageNumber="287" phylum="Cnidaria" rank="species" species="heterosceles">
<emphasis box="[151,398,1523,1544]" italics="true" pageId="51" pageNumber="287">Staurodiscus heterosceles</emphasis>
<bibRefCitation author="Haeckel" box="[404,594,1523,1545]" firstAuthor="Haeckel" page="146" pageId="51" pageNumber="287" pagination="1 - 360" refId="ref68751" refString="Haeckel E. 1879. Das System der Medusen. Erster Teil einer Monographie der Medusen. Denkschriften der Medicinisch- Naturwissenschaftlichen Gesellschaft zu Jena 1: I-XX, 1 - 360, 20 pls." type="book chapter" year="1879">Haeckel, 1879: 146</bibRefCitation>
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. 
<emphasis bold="true" box="[603,692,1523,1544]" pageId="51" pageNumber="287">syn. nov.</emphasis>
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<paragraph blockId="51.[151,757,1568,2077]" pageId="51" pageNumber="287">
<emphasis bold="true" box="[151,379,1568,1592]" pageId="51" pageNumber="287">Examined material:</emphasis>
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; 
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; 
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<collectingDate pageId="51" pageNumber="287" value="2018-11-15">15-NOV- 2018</collectingDate>
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; size 
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; preserved in ethanol for DNA extraction; 16S sequence 
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. 
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– 
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; 
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; size 
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; preserved in ethanol for DNA extraction; 16S sequence 
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. 
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– 
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; 
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; 
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<collectingDate box="[458,617,1730,1754]" pageId="51" pageNumber="287" value="2020-05-30">30-MAY-2020</collectingDate>
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; preserved in ethanol for DNA extraction; 16S sequence identical 
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. 
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– 
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; 
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; 
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; preserved in ethanol for DNA extraction; 16S sequence identical to 
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. 
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– 
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; 
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; 
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; size 
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; part preserved in formalin and deposited as UF-014059, small part preserved in ethanol for DNA extraction; 16S sequence identical to 
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. 
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– 
<date box="[596,752,1989,2013]" pageId="51" pageNumber="287" value="2017-12-11">
<collectingDate box="[596,752,1989,2013]" pageId="51" pageNumber="287" value="2017-12-11">11-DEC-2017</collectingDate>
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photographed; 
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, gonads not yet developed, not collected.
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</paragraph>
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<subSubSection pageId="51" pageNumber="287" type="discussion">
<paragraph blockId="51.[832,1437,257,969]" pageId="51" pageNumber="287">
<emphasis bold="true" box="[832,995,257,281]" pageId="51" pageNumber="287">Observations:</emphasis>
Bell hemispherical, 
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when mature, apical jelly about 1/3 of total height. Manubrium composed of central, cylindrical part and four basal, perradial extensions each with two lateral outgrowths in about middle of bell. Central part of manubrium tubular, with cruciform mouth opening drawn into four lips, margin slightly undulated. Basal, perradial extensions of stomach reaching almost to circular canal and thus length of radial canals to circular canal is short (1/4); stomach outgrowths relatively thick, giving off lateral diverticula in about the middle, these paired outgrowths opposite or not, oblique to radial axis or slightly curved, towards ends somewhat irregularly folded or undulating, not connecting to circular canal. Gonads in walls of stomach outgrowths. Mature animals with 23-30 tentacles and 1-3 cordyli between each pair of tentacles. Young animal with four perradial tentacles only and 7-9 cordyli per quadrant (
<figureCitation box="[1215,1316,789,813]" captionStart="Fig" captionStartId="52.[151,186,1998,2020]" captionTargetBox="[151,1436,648,1932]" captionTargetId="figure-165@52.[151,1436,648,1932]" captionTargetPageId="52" captionText="Fig. 32. Staurodiscus tetrastaurus. (A-C) Young animal observed 11-DEC-2017. (D-E) BFLA3831, size 7 mm. Some amphipods are associated with the animal. (F) BFLA4440, size 10 mm, bell margin seen from adaxial side, ocelli are barely visible. (G) BFLA4440, oral view, note variation of position of lateral diverticula." figureDoi="http://doi.org/10.5281/zenodo.5640032" httpUri="https://zenodo.org/record/5640032/files/figure.png" pageId="51" pageNumber="287">Fig. 32A</figureCitation>
). Tentacles with conical bulbs. Small, inconspicuous ocelli present on adaxial side of tentacles bulbs and attachment sites of cordyli (
<figureCitation box="[1020,1121,882,907]" captionStart="Fig" captionStartId="52.[151,186,1998,2020]" captionTargetBox="[151,1436,648,1932]" captionTargetId="figure-165@52.[151,1436,648,1932]" captionTargetPageId="52" captionText="Fig. 32. Staurodiscus tetrastaurus. (A-C) Young animal observed 11-DEC-2017. (D-E) BFLA3831, size 7 mm. Some amphipods are associated with the animal. (F) BFLA4440, size 10 mm, bell margin seen from adaxial side, ocelli are barely visible. (G) BFLA4440, oral view, note variation of position of lateral diverticula." figureDoi="http://doi.org/10.5281/zenodo.5640032" httpUri="https://zenodo.org/record/5640032/files/figure.png" pageId="51" pageNumber="287">Fig. 32F</figureCitation>
). Animal colourless or with greenish tint. Amphipods appear often to associate with the medusa (
<figureCitation box="[967,1047,945,969]" captionStart="Fig" captionStartId="52.[151,186,1998,2020]" captionTargetBox="[151,1436,648,1932]" captionTargetId="figure-165@52.[151,1436,648,1932]" captionTargetPageId="52" captionText="Fig. 32. Staurodiscus tetrastaurus. (A-C) Young animal observed 11-DEC-2017. (D-E) BFLA3831, size 7 mm. Some amphipods are associated with the animal. (F) BFLA4440, size 10 mm, bell margin seen from adaxial side, ocelli are barely visible. (G) BFLA4440, oral view, note variation of position of lateral diverticula." figureDoi="http://doi.org/10.5281/zenodo.5640032" httpUri="https://zenodo.org/record/5640032/files/figure.png" pageId="51" pageNumber="287">Fig. 32</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection pageId="51" pageNumber="287" type="description">
<paragraph blockId="51.[832,1437,991,1203]" pageId="51" pageNumber="287">
<emphasis bold="true" box="[832,945,991,1015]" pageId="51" pageNumber="287">16S Data:</emphasis>
The five obtained 16S sequences represented 2 haplotypes differing only one position (an insertion/ deletion). In the maximum likelihood tree (
<figureCitation box="[1297,1375,1054,1078]" captionStart="Fig" captionStartId="45.[153,188,1968,1990]" captionTargetBox="[199,1370,677,1924]" captionTargetId="graphics-159@45.[199,969,686,1866]" captionTargetPageId="45" captionText="Fig. 28. 16S maximum likelihood phylogenetictree of Laodiceidae,Hebellidae,and Orchistomatidae,obtainedwith PhyML (GTR+G+I model) and based on about 600 bp positions of the mitochondrial 16S gene. Node-support values are bootstrap values of 100 pseudoreplicates (shown only if&gt; 70%).All sequence labels start with the GenBank numbers (except for identical haplotypes) permitting the retrieval of more information. Red labels are new sequences from this study." figureDoi="http://doi.org/10.5281/zenodo.5640020" httpUri="https://zenodo.org/record/5640020/files/figure.png" pageId="51" pageNumber="287">Fig. 28</figureCitation>
) they are well separated from the other 
<taxonomicName box="[1212,1436,1085,1109]" class="Hydrozoa" family="Hebellidae" genus="Staurodiscus" kingdom="Animalia" order="Leptothecata" pageId="51" pageNumber="287" phylum="Cnidaria" rank="species" species="undetermined">
<emphasis box="[1212,1349,1085,1109]" italics="true" pageId="51" pageNumber="287">Staurodiscus</emphasis>
species
</taxonomicName>
and also 
<taxonomicName authorityName="Haeckel" authorityYear="1879" box="[933,1065,1116,1140]" class="Hydrozoa" family="Laodiceidae" genus="Melicertissa" kingdom="Animalia" order="Leptothecata" pageId="51" pageNumber="287" phylum="Cnidaria" rank="genus">
<emphasis box="[933,1065,1116,1140]" italics="true" pageId="51" pageNumber="287">Melicertissa</emphasis>
</taxonomicName>
of this study. It is part of a well supported clade comprising the 
<taxonomicName box="[1182,1405,1148,1172]" class="Hydrozoa" family="Hebellidae" genus="Staurodiscus" kingdom="Animalia" order="Leptothecata" pageId="51" pageNumber="287" phylum="Cnidaria" rank="species" species="undetermined">
<emphasis box="[1182,1319,1148,1172]" italics="true" pageId="51" pageNumber="287">Staurodiscus</emphasis>
species
</taxonomicName>
of this study and some 
<taxonomicName authorityName="Fraser" authorityYear="1912" box="[1049,1168,1179,1203]" class="Hydrozoa" family="Hebellidae" kingdom="Animalia" order="Leptothecata" pageId="51" pageNumber="287" phylum="Cnidaria" rank="family">Hebellidae</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection pageId="51" pageNumber="287" type="distribution">
<paragraph blockId="51.[832,1437,1225,1406]" pageId="51" pageNumber="287">
<emphasis bold="true" box="[832,984,1225,1249]" pageId="51" pageNumber="287">Distribution:</emphasis>
Canary Islands, Florida, Caribbean Sea, coast of 
<collectingCountry box="[984,1128,1256,1280]" name="Mozambique" pageId="51" pageNumber="287">Mozambique</collectingCountry>
, coasts of 
<collectingCountry box="[1249,1307,1256,1280]" name="India" pageId="51" pageNumber="287">India</collectingCountry>
, 
<collectingCountry box="[1321,1432,1256,1280]" name="Singapore" pageId="51" pageNumber="287">Singapore</collectingCountry>
, 
<collectingCountry box="[832,911,1288,1312]" name="Taiwan" pageId="51" pageNumber="287">Taiwan</collectingCountry>
Strait, Bismarck Sea (
<bibRefCitation author="Kramp" box="[1188,1355,1288,1312]" firstAuthor="Kramp" pageId="51" pageNumber="287" pagination="1 - 283" refId="ref69812" refString="Kramp P. L. 1959 a. The Hydromedusae of the Atlantic Ocean and adjacent waters. Dana Report 46: 1 - 283." type="book chapter" year="1959" yearSuffix="a">Kramp, 1959a</bibRefCitation>
, 
<bibRefCitation author="Kramp" box="[1375,1432,1288,1312]" firstAuthor="Kramp" pageId="51" pageNumber="287" pagination="1 - 200" refId="ref69950" refString="Kramp P. L. 1968. The hydromedusae of the Pacific and Indian oceans. Sections II and III. Dana Report 72: 1 - 200." type="book chapter" year="1968">1968</bibRefCitation>
; 
<bibRefCitation author="Bouillon" box="[832,1013,1319,1343]" firstAuthor="Bouillon" pageId="51" pageNumber="287" pagination="117 - 172" refId="ref65593" refString="Bouillon J. 1978 a. Hydromeduses de l'Archipel des Sechelles et du Mozambique. Revue de Zoologie Africaine 92 (1): 117 - 172." type="journal article" year="1978" yearSuffix="a">Bouillon 1978a</bibRefCitation>
, 
<bibRefCitation author="Bouillon" box="[1036,1107,1319,1343]" firstAuthor="Bouillon" pageId="51" pageNumber="287" pagination="25 - 112" refId="ref65793" refString="Bouillon J. 1984 b. Hydromeduses de la Mer de Bismarck (Papouasie Nouvelle-Guinee). Partie IV: Leptomedusae (Hydrozoa - Cnidaria). Indo-Malayan Zoology 1 (1): 25 - 112." type="journal article" year="1984" yearSuffix="b">1984b</bibRefCitation>
; Navas-Pereira &amp; Vannuci, 1991; 
<bibRefCitation author="Zhang" box="[903,1045,1350,1374]" firstAuthor="Zhang" pageId="51" pageNumber="287" pagination="76 - 82" refId="ref77079" refString="Zhang J. 1999. Hydromedusae and Siphonophora in western waters of Taiwan Island during winter and spring. Journal of Oceanography in Taiwan Strait 18 (1): 76 - 82." type="journal article" year="1999">Zhang, 1999</bibRefCitation>
; 
<bibRefCitation author="Segura-Puertas" box="[1059,1363,1350,1374]" etAl="et al." firstAuthor="Segura-Puertas" pageId="51" pageNumber="287" pagination="1 - 15" refId="ref74885" refString="Segura-Puertas L., Suarez Morales E., Celis L. 2003. A checklist of the medusae (Hydrozoa, Scyphozoa and Cubozoa) of Mexico. Zootaxa 194: 1 - 15." type="journal article" year="2003">
Segura-Puertas 
<emphasis box="[1231,1288,1350,1374]" italics="true" pageId="51" pageNumber="287">et al.</emphasis>
, 2003
</bibRefCitation>
). 
<typeStatus box="[1383,1436,1351,1375]" pageId="51" pageNumber="287">Type</typeStatus>
locality: Atlantic Ocean, Canary Islands, near Lanzarote.
</paragraph>
</subSubSection>
<subSubSection lastPageId="52" lastPageNumber="288" pageId="51" pageNumber="287" type="discussion">
<paragraph blockId="51.[832,1437,1428,2077]" pageId="51" pageNumber="287">
<emphasis bold="true" box="[832,946,1428,1452]" pageId="51" pageNumber="287">Remarks:</emphasis>
<bibRefCitation author="Haeckel" box="[967,1150,1428,1452]" firstAuthor="Haeckel" pageId="51" pageNumber="287" pagination="1 - 360" refId="ref68751" refString="Haeckel E. 1879. Das System der Medusen. Erster Teil einer Monographie der Medusen. Denkschriften der Medicinisch- Naturwissenschaftlichen Gesellschaft zu Jena 1: I-XX, 1 - 360, 20 pls." type="book chapter" year="1879">Haeckel (1879)</bibRefCitation>
introduced two nominal 
<taxonomicName box="[890,1138,1459,1483]" class="Hydrozoa" family="Hebellidae" genus="Staurodiscus" kingdom="Animalia" order="Leptothecata" pageId="51" pageNumber="287" phylum="Cnidaria" rank="species" species="undetermined">
<emphasis box="[890,1031,1459,1483]" italics="true" pageId="51" pageNumber="287">Staurodiscus</emphasis>
species
</taxonomicName>
, 
<taxonomicName authorityName="Haeckel" authorityYear="1879" box="[1168,1372,1460,1483]" class="Hydrozoa" family="Hebellidae" genus="Staurodiscus" kingdom="Animalia" order="Leptothecata" pageId="51" pageNumber="287" phylum="Cnidaria" rank="species" species="tetrastaurus">
<emphasis box="[1168,1372,1460,1483]" italics="true" pageId="51" pageNumber="287">S. tetrastaurus</emphasis>
</taxonomicName>
and 
<taxonomicName authorityName="Haeckel" authorityYear="1879" box="[832,997,1490,1514]" class="Hydrozoa" family="Hebellidae" genus="Staurodiscus" kingdom="Animalia" order="Leptothecata" pageId="51" pageNumber="287" phylum="Cnidaria" rank="species" species="heterosceles">
<emphasis box="[832,997,1490,1514]" italics="true" pageId="51" pageNumber="287">S. heterosceles</emphasis>
</taxonomicName>
, both collected at the same time and near Lanzarote Island. The former had 8-16 tentacles, 48 cordyli, and a bell diameter of 
<quantity box="[1229,1321,1553,1577]" metricMagnitude="-3" metricUnit="m" metricValue="5.0" metricValueMax="6.0" metricValueMin="4.0" pageId="51" pageNumber="287" unit="mm" value="5.0" valueMax="6.0" valueMin="4.0">4-6 mm</quantity>
, the latter 8-32 tentacles, 40-80 cordyli, and a bell diameter of 
<quantity box="[832,918,1615,1639]" metricMagnitude="-3" metricUnit="m" metricValue="7.0" metricValueMax="8.0" metricValueMin="6.0" pageId="51" pageNumber="287" unit="mm" value="7.0" valueMax="8.0" valueMin="6.0">6-8 mm</quantity>
[note that the figures in 
<bibRefCitation author="Kramp" box="[1199,1359,1615,1639]" firstAuthor="Kramp" pageId="51" pageNumber="287" pagination="1 - 283" refId="ref69812" refString="Kramp P. L. 1959 a. The Hydromedusae of the Atlantic Ocean and adjacent waters. Dana Report 46: 1 - 283." type="book chapter" year="1959" yearSuffix="a">Kramp (1959a</bibRefCitation>
, 
<bibRefCitation author="Kramp" box="[1374,1429,1615,1639]" firstAuthor="Kramp" pageId="51" pageNumber="287" pagination="1 - 200" refId="ref69950" refString="Kramp P. L. 1968. The hydromedusae of the Pacific and Indian oceans. Sections II and III. Dana Report 72: 1 - 200." type="book chapter" year="1968">1968</bibRefCitation>
) depict a juvenile 
<taxonomicName authorityName="Haeckel" authorityYear="1879" box="[1037,1201,1647,1670]" class="Hydrozoa" family="Hebellidae" genus="Staurodiscus" kingdom="Animalia" order="Leptothecata" pageId="51" pageNumber="287" phylum="Cnidaria" rank="species" species="tetrastaurus">
<emphasis box="[1037,1201,1647,1670]" italics="true" pageId="51" pageNumber="287">S. tetrastaurus</emphasis>
</taxonomicName>
with four tentacles, compare 
<figureCitation box="[932,1033,1678,1702]" captionStart="Fig" captionStartId="52.[151,186,1998,2020]" captionTargetBox="[151,1436,648,1932]" captionTargetId="figure-165@52.[151,1436,648,1932]" captionTargetPageId="52" captionText="Fig. 32. Staurodiscus tetrastaurus. (A-C) Young animal observed 11-DEC-2017. (D-E) BFLA3831, size 7 mm. Some amphipods are associated with the animal. (F) BFLA4440, size 10 mm, bell margin seen from adaxial side, ocelli are barely visible. (G) BFLA4440, oral view, note variation of position of lateral diverticula." figureDoi="http://doi.org/10.5281/zenodo.5640032" httpUri="https://zenodo.org/record/5640032/files/figure.png" pageId="51" pageNumber="287">Fig. 32A</figureCitation>
]. Additionally, Haeckel distinguished both nominal species based on the branching pattern of the radial canals (= actually basal extensions of the manubrium, comp. 
<figureCitation box="[1058,1244,1772,1796]" captionStart="Fig" captionStartId="52.[151,186,1998,2020]" captionTargetBox="[151,1436,648,1932]" captionTargetId="figure-165@52.[151,1436,648,1932]" captionTargetPageId="52" captionText="Fig. 32. Staurodiscus tetrastaurus. (A-C) Young animal observed 11-DEC-2017. (D-E) BFLA3831, size 7 mm. Some amphipods are associated with the animal. (F) BFLA4440, size 10 mm, bell margin seen from adaxial side, ocelli are barely visible. (G) BFLA4440, oral view, note variation of position of lateral diverticula." figureDoi="http://doi.org/10.5281/zenodo.5640032" httpUri="https://zenodo.org/record/5640032/files/figure.png" pageId="51" pageNumber="287">Fig. 32A, E, G</figureCitation>
): 
<taxonomicName authorityName="Haeckel" authorityYear="1879" box="[1271,1436,1772,1795]" class="Hydrozoa" family="Hebellidae" genus="Staurodiscus" kingdom="Animalia" order="Leptothecata" pageId="51" pageNumber="287" phylum="Cnidaria" rank="species" species="tetrastaurus">
<emphasis box="[1271,1436,1772,1795]" italics="true" pageId="51" pageNumber="287">S. tetrastaurus</emphasis>
</taxonomicName>
has opposite pairs, while they were not opposite in 
<emphasis box="[873,1054,1834,1858]" italics="true" pageId="51" pageNumber="287">
<taxonomicName authorityName="Haeckel" authorityYear="1879" box="[873,1050,1834,1858]" class="Hydrozoa" family="Hebellidae" genus="Staurodiscus" kingdom="Animalia" order="Leptothecata" pageId="51" pageNumber="287" phylum="Cnidaria" rank="species" species="heterosceles">S. heterosceles</taxonomicName>
.
</emphasis>
<bibRefCitation author="Haeckel" box="[1073,1255,1834,1858]" firstAuthor="Haeckel" pageId="51" pageNumber="287" pagination="1 - 360" refId="ref68751" refString="Haeckel E. 1879. Das System der Medusen. Erster Teil einer Monographie der Medusen. Denkschriften der Medicinisch- Naturwissenschaftlichen Gesellschaft zu Jena 1: I-XX, 1 - 360, 20 pls." type="book chapter" year="1879">Haeckel (1879)</bibRefCitation>
also observed intermediate forms and explicitly expressed doubts that they are “ 
<emphasis box="[945,1086,1897,1921]" italics="true" pageId="51" pageNumber="287">bona species</emphasis>
”, but nevertheless named both forms [Haeckel used an artificial, typological species concept and did not distinguish biological species, comp. comments in 
<bibRefCitation author="Haeckel" box="[1055,1215,1990,2015]" firstAuthor="Haeckel" pageId="51" pageNumber="287" pagination="1 - 360" refId="ref68751" refString="Haeckel E. 1879. Das System der Medusen. Erster Teil einer Monographie der Medusen. Denkschriften der Medicinisch- Naturwissenschaftlichen Gesellschaft zu Jena 1: I-XX, 1 - 360, 20 pls." type="book chapter" year="1879">Haeckel, 1879</bibRefCitation>
: XXV, 109]. 
<bibRefCitation author="Kramp" firstAuthor="Kramp" pageId="51" pageNumber="287" pagination="305 - 366" refId="ref69901" refString="Kramp P. L. 1962. Medusae of Vietnam. Videnskabelige Meddelelser fra Dansk naturhistorisk Forening i KObenhavn 124: 305 - 366." type="journal article" year="1962">Kramp (1962</bibRefCitation>
, 
<bibRefCitation author="Kramp" box="[908,963,2022,2046]" firstAuthor="Kramp" pageId="51" pageNumber="287" pagination="1 - 200" refId="ref69950" refString="Kramp P. L. 1968. The hydromedusae of the Pacific and Indian oceans. Sections II and III. Dana Report 72: 1 - 200." type="book chapter" year="1968">1968</bibRefCitation>
) too thought that both names refer likely to the same species.
</paragraph>
<paragraph blockId="52.[151,756,257,596]" pageId="52" pageNumber="288">
One of our specimen (BFLA3831) was also intermediate as it had the size and tentacle numbers of 
<emphasis box="[592,756,289,313]" italics="true" pageId="52" pageNumber="288">
<taxonomicName authorityName="Haeckel" authorityYear="1879" box="[592,751,289,313]" class="Hydrozoa" family="Hebellidae" genus="Staurodiscus" kingdom="Animalia" order="Leptothecata" pageId="52" pageNumber="288" phylum="Cnidaria" rank="species" species="heterosceles">S. heterosceles</taxonomicName>
,
</emphasis>
but the outgrowths of the stomach had opposite sidebranches (
<figureCitation box="[265,366,352,376]" captionStart="Fig" captionStartId="52.[151,186,1998,2020]" captionTargetBox="[151,1436,648,1932]" captionTargetId="figure-165@52.[151,1436,648,1932]" captionTargetPageId="52" captionText="Fig. 32. Staurodiscus tetrastaurus. (A-C) Young animal observed 11-DEC-2017. (D-E) BFLA3831, size 7 mm. Some amphipods are associated with the animal. (F) BFLA4440, size 10 mm, bell margin seen from adaxial side, ocelli are barely visible. (G) BFLA4440, oral view, note variation of position of lateral diverticula." figureDoi="http://doi.org/10.5281/zenodo.5640032" httpUri="https://zenodo.org/record/5640032/files/figure.png" pageId="52" pageNumber="288">Fig. 32E</figureCitation>
). Others (
<figureCitation box="[478,580,352,376]" captionStart="Fig" captionStartId="52.[151,186,1998,2020]" captionTargetBox="[151,1436,648,1932]" captionTargetId="figure-165@52.[151,1436,648,1932]" captionTargetPageId="52" captionText="Fig. 32. Staurodiscus tetrastaurus. (A-C) Young animal observed 11-DEC-2017. (D-E) BFLA3831, size 7 mm. Some amphipods are associated with the animal. (F) BFLA4440, size 10 mm, bell margin seen from adaxial side, ocelli are barely visible. (G) BFLA4440, oral view, note variation of position of lateral diverticula." figureDoi="http://doi.org/10.5281/zenodo.5640032" httpUri="https://zenodo.org/record/5640032/files/figure.png" pageId="52" pageNumber="288">Fig. 32G</figureCitation>
) had a variable branching pattern. The observed juvenile specimen (
<figureCitation box="[159,258,415,439]" captionStart="Fig" captionStartId="52.[151,186,1998,2020]" captionTargetBox="[151,1436,648,1932]" captionTargetId="figure-165@52.[151,1436,648,1932]" captionTargetPageId="52" captionText="Fig. 32. Staurodiscus tetrastaurus. (A-C) Young animal observed 11-DEC-2017. (D-E) BFLA3831, size 7 mm. Some amphipods are associated with the animal. (F) BFLA4440, size 10 mm, bell margin seen from adaxial side, ocelli are barely visible. (G) BFLA4440, oral view, note variation of position of lateral diverticula." figureDoi="http://doi.org/10.5281/zenodo.5640032" httpUri="https://zenodo.org/record/5640032/files/figure.png" pageId="52" pageNumber="288">Fig. 32A</figureCitation>
) had three opposite and one alternate pairs of side-branches. It is thus quite clear that 
<taxonomicName authorityName="Haeckel" authorityYear="1879" box="[597,756,447,470]" class="Hydrozoa" family="Hebellidae" genus="Staurodiscus" kingdom="Animalia" order="Leptothecata" pageId="52" pageNumber="288" phylum="Cnidaria" rank="species" species="tetrastaurus">
<emphasis box="[597,756,447,470]" italics="true" pageId="52" pageNumber="288">S. tetrastaurus</emphasis>
</taxonomicName>
and 
<taxonomicName authorityName="Haeckel" authorityYear="1879" box="[203,368,478,502]" class="Hydrozoa" family="Hebellidae" genus="Staurodiscus" kingdom="Animalia" order="Leptothecata" pageId="52" pageNumber="288" phylum="Cnidaria" rank="species" species="heterosceles">
<emphasis box="[203,368,478,502]" italics="true" pageId="52" pageNumber="288">S. heterosceles</emphasis>
</taxonomicName>
must be regarded as conspecific, 
<taxonomicName authorityName="Haeckel" authorityYear="1879" box="[151,307,509,533]" class="Hydrozoa" family="Hebellidae" genus="Staurodiscus" kingdom="Animalia" order="Leptothecata" pageId="52" pageNumber="288" phylum="Cnidaria" rank="species" species="tetrastaurus">
<emphasis box="[151,307,509,533]" italics="true" pageId="52" pageNumber="288">S. tetrastaurus</emphasis>
</taxonomicName>
being the valid name. Medusae identified as 
<taxonomicName authorityName="Haeckel" authorityYear="1879" box="[182,457,540,564]" class="Hydrozoa" family="Hebellidae" genus="Staurodiscus" kingdom="Animalia" order="Leptothecata" pageId="52" pageNumber="288" phylum="Cnidaria" rank="species" species="heterosceles">
<emphasis box="[182,457,540,564]" italics="true" pageId="52" pageNumber="288">Staurodiscus heterosceles</emphasis>
</taxonomicName>
have – to our knowledge – not been recorded anymore since Haeckel’s time.
</paragraph>
<paragraph blockId="52.[832,1437,257,502]" pageId="52" pageNumber="288">
<bibRefCitation author="Andrade &amp; Migotto" box="[832,1134,257,281]" firstAuthor="Andrade" pageId="52" pageNumber="288" pagination="35 - 36" refId="ref64532" refString="Andrade L., Migotto A. 1997. Is there a link between Hebella Hydroids (Hydrozoa, Lafoeidae) and Staurodiscus medusae (Hydrozoa, Laodiceidae)? Resumos expandidos do VII COLACMAR Congresso Latino-Americano sobre Ciencias do Mar, I: 35 - 36." type="journal article" year="1997">Andrade &amp; Migotto (1997)</bibRefCitation>
reared the initial stages of medusae released from a hydroid referable to 
<taxonomicName authority="(Bale, 1888)" baseAuthorityName="Bale" baseAuthorityYear="1888" class="Hydrozoa" family="Hebellidae" genus="Hebella" kingdom="Animalia" order="Leptothecata" pageId="52" pageNumber="288" phylum="Cnidaria" rank="species" species="scandens">
<emphasis italics="true" pageId="52" pageNumber="288">Hebella scandens</emphasis>
(
<bibRefCitation author="Bale" box="[949,1073,320,344]" firstAuthor="Bale" pageId="52" pageNumber="288" pagination="745 - 799" refId="ref64631" refString="Bale W. M. 1888. On some new and rare Hydroida in the Australian Museum collections. Proceedings of the Linnean Society of New South Wales ser. 2 volume 3: 745 - 799, pls 12 - 21." type="journal article" year="1888">Bale, 1888</bibRefCitation>
)
</taxonomicName>
and found that they resembled our 
<emphasis box="[878,1046,352,375]" italics="true" pageId="52" pageNumber="288">
<taxonomicName authorityName="Haeckel" authorityYear="1879" box="[878,1041,352,375]" class="Hydrozoa" family="Hebellidae" genus="Staurodiscus" kingdom="Animalia" order="Leptothecata" pageId="52" pageNumber="288" phylum="Cnidaria" rank="species" species="tetrastaurus">S. tetrastaurus</taxonomicName>
.
</emphasis>
Our 16S phylogeny (
<figureCitation box="[1293,1375,352,376]" captionStart="Fig" captionStartId="45.[153,188,1968,1990]" captionTargetBox="[199,1370,677,1924]" captionTargetId="graphics-159@45.[199,969,686,1866]" captionTargetPageId="45" captionText="Fig. 28. 16S maximum likelihood phylogenetictree of Laodiceidae,Hebellidae,and Orchistomatidae,obtainedwith PhyML (GTR+G+I model) and based on about 600 bp positions of the mitochondrial 16S gene. Node-support values are bootstrap values of 100 pseudoreplicates (shown only if&gt; 70%).All sequence labels start with the GenBank numbers (except for identical haplotypes) permitting the retrieval of more information. Red labels are new sequences from this study." figureDoi="http://doi.org/10.5281/zenodo.5640020" httpUri="https://zenodo.org/record/5640020/files/figure.png" pageId="52" pageNumber="288">Fig. 28</figureCitation>
) also suggests a more close relationship to 
<taxonomicName authorityName="Fraser" authorityYear="1912" box="[1263,1379,383,407]" class="Hydrozoa" family="Hebellidae" kingdom="Animalia" order="Leptothecata" pageId="52" pageNumber="288" phylum="Cnidaria" rank="family">Hebellidae</taxonomicName>
than to 
<taxonomicName authorityName="L. Agassiz" authorityYear="1862" box="[866,1000,415,439]" class="Hydrozoa" family="Laodiceidae" kingdom="Animalia" order="Leptothecata" pageId="52" pageNumber="288" phylum="Cnidaria" rank="family">Laodiceidae</taxonomicName>
. 
<taxonomicName authorityName="Haeckel" authorityYear="1879" box="[1018,1298,415,439]" class="Hydrozoa" family="Hebellidae" genus="Staurodiscus" kingdom="Animalia" order="Leptothecata" pageId="52" pageNumber="288" phylum="Cnidaria" rank="species" species="tetrastaurus">
<emphasis box="[1018,1298,415,439]" italics="true" pageId="52" pageNumber="288">Staurodiscus tetrastaurus</emphasis>
</taxonomicName>
is the 
<typeStatus box="[1390,1436,415,439]" pageId="52" pageNumber="288">type</typeStatus>
species of the genus, thus decisive to which family the genus belongs.
</paragraph>
</subSubSection>
</treatment>
</document>