treatments-xml/data/03/E8/87/03E887962270B04E2565EF19FB01F907.xml
2024-06-21 12:22:17 +02:00

1913 lines
292 KiB
XML
Raw Blame History

This file contains ambiguous Unicode characters

This file contains Unicode characters that might be confused with other characters. If you think that this is intentional, you can safely ignore this warning. Use the Escape button to reveal them.

<document id="02C7936BAC7764FD30ADC04EE94657BE" ID-DOI="10.1080/14772019.2017.1371257" ID-ISSN="1478-0941" ID-PMC="PMC6130837" ID-PubMed="30210265" ID-Zenodo-Dep="10883262" ID-ZooBank="EDD6E170-CA64-4FFB-8DD1-AED2D61D5504" IM.bibliography_approvedBy="diego" IM.illustrations_approvedBy="diego" IM.materialsCitations_approvedBy="diego" IM.metadata_approvedBy="felipe" IM.tables_approvedBy="diego" IM.taxonomicNames_approvedBy="felipe" IM.treatments_approvedBy="diego" checkinTime="1711483590210" checkinUser="felipe" docAuthor="Marram ̀, Giuseppe, Claeso, Kerin M., Carneval, Giorgio &amp; Kriwe, Jurgen" docDate="2018" docId="03E887962270B04E2565EF19FB01F907" docLanguage="en" docName="JSystPalaeontol.16.14.1189-1219.pdf" docOrigin="Journal of Systematic Palaeontology 16 (14)" docSource="http://dx.doi.org/10.1080/14772019.2017.1371257" docStyle="DocumentStyle:6400638016C1F4AE0D0B6502E032D571.1:JSystPalaeontol.2016-.journal_article.closed" docStyleId="6400638016C1F4AE0D0B6502E032D571" docStyleName="JSystPalaeontol.2016-.journal_article.closed" docStyleVersion="1" docTitle="Titanonarke molini" docType="treatment" docVersion="4" lastPageNumber="1205" masterDocId="FFD1FFEE2275B05F2472E94CFFD9FFCC" masterDocTitle="Revision of Eocene electric rays (Torpediniformes, Batomorphii) from the Bolca Konservat-Lagerst atte, Italy, reveals the first fossil embryo in situ in marine batoids and provides new insights into the origin of trophic novelties in coral reef fishes" masterLastPageNumber="1219" masterPageNumber="1189" pageNumber="1193" updateTime="1712145816225" updateUser="diego">
<mods:mods id="AE674EDC69653DD9931991AEF235555C" xmlns:mods="http://www.loc.gov/mods/v3">
<mods:titleInfo id="0BD9816ABECEC6CC1A64EB7C573E98F7">
<mods:title id="2166314C085820FFE2BFED3F921B9BE5">Revision of Eocene electric rays (Torpediniformes, Batomorphii) from the Bolca Konservat-Lagerst atte, Italy, reveals the first fossil embryo in situ in marine batoids and provides new insights into the origin of trophic novelties in coral reef fishes</mods:title>
</mods:titleInfo>
<mods:name id="A79272D6CA54F69860AF6FEE0B09627E" type="personal">
<mods:role id="EBB6182D6635BA7884963FEDEB5C0007">
<mods:roleTerm id="2FFCCB0C4CB82D9F785264B6F2B31A9B">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="BFEBC980F20652EA36B61B4D5CB630D6">Marram ̀, Giuseppe</mods:namePart>
<mods:affiliation id="EBF7F8C0552E9E13ABF006B2066D0B62">University of Vienna, Department of Palaeontology, Althanstrasse 14, 1090, Vienna;</mods:affiliation>
</mods:name>
<mods:name id="41AA8374E1D11738251F7FCE0B9034A9" type="personal">
<mods:role id="A16FC322CFE1604794A7DCB20569894C">
<mods:roleTerm id="45FFC695EAB7060F980EFD0DEE21F42D">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="6D0D6161D5FD4AE23B67683A6C903112">Claeso, Kerin M.</mods:namePart>
</mods:name>
<mods:name id="13A652EA7C22A4FA2D94CF4DF5F20149" type="personal">
<mods:role id="245CBD924F4B7F1777F8D3F0C40D95CC">
<mods:roleTerm id="8847D8E96B0F6600D34D8B1417B112C0">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="C75C8402725EB1E462B950A8A0A5F85B">Carneval, Giorgio</mods:namePart>
</mods:name>
<mods:name id="47AC1F5B3A66E6C0F56027EB2D56ED93" type="personal">
<mods:role id="C691DFE3F05B7B4CEE0D1213F5EEE10D">
<mods:roleTerm id="59DF9F1D6D69720F97A57B42EFA0B534">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="22830662D94C349EFC831C1D964FB488">Kriwe, Jurgen</mods:namePart>
</mods:name>
<mods:typeOfResource id="DD4AD5C791346D244E01747783E0DB6A">text</mods:typeOfResource>
<mods:relatedItem id="A9ABF12186102EBC88AC51DEFDD99D02" type="host">
<mods:titleInfo id="B1C2D037EA55993B4900AB025B20290D">
<mods:title id="D9E5E91314D94966B3A7773FC4F960AD">Journal of Systematic Palaeontology</mods:title>
</mods:titleInfo>
<mods:part id="4186AE02F37152C8EB920FC85F999878">
<mods:date id="2A3FC2BE4541348BCF63163857F1FE75">2018</mods:date>
<mods:detail id="CD381EC31A0A37249F0BD28BAD7BFAC2" type="pubDate">
<mods:number id="C253ECDEC0EDE49BB99178B9D03329FC">2017-09-21</mods:number>
</mods:detail>
<mods:detail id="DFC31A58664AD9C7DA52CC86540FA0C2" type="volume">
<mods:number id="E402B2721DD102DD1C4F28310C89FFD8">16</mods:number>
</mods:detail>
<mods:detail id="CF3D54F8FAEA71FF91400124DC2780CB" type="issue">
<mods:number id="8CB4512626F962D06984A3BF650C8B1A">14</mods:number>
</mods:detail>
<mods:extent id="6FB030329A2FAA2C813605E690EA923A" unit="page">
<mods:start id="12F9C52338EFC584A5EC3E0097FEF514">1189</mods:start>
<mods:end id="5CEB29D49CF3F9850FEC6263C7E657B8">1219</mods:end>
</mods:extent>
</mods:part>
</mods:relatedItem>
<mods:location id="C5B16B48BEB5740458E3538ACC68786A">
<mods:url id="802F09AF08CD315CB16A3972A589D150">http://dx.doi.org/10.1080/14772019.2017.1371257</mods:url>
</mods:location>
<mods:classification id="D19BDABF1C4C79B735D84B04C794705D">journal article</mods:classification>
<mods:identifier id="B6E54F43B23F26E70A3ECC741ADC4F3E" type="DOI">10.1080/14772019.2017.1371257</mods:identifier>
<mods:identifier id="F25D96FF93DA87D1547D0FE3B25D1D65" type="ISSN">1478-0941</mods:identifier>
<mods:identifier id="BF3DA04F9808A28E42E0A9C6199CDE86" type="PMC">PMC6130837</mods:identifier>
<mods:identifier id="B4027B26F257CAC00CA3F8CD0B4CF806" type="PubMed">30210265</mods:identifier>
<mods:identifier id="CFC23942E47AFDBE414E97543BA8484D" type="Zenodo-Dep">10883262</mods:identifier>
<mods:identifier id="49E3FFC610DB3A9FCAD44A9EA28EAAEC" type="ZooBank">EDD6E170-CA64-4FFB-8DD1-AED2D61D5504</mods:identifier>
</mods:mods>
<treatment id="03E887962270B04E2565EF19FB01F907" LSID="urn:lsid:plazi:treatment:03E887962270B04E2565EF19FB01F907" httpUri="http://treatment.plazi.org/id/03E887962270B04E2565EF19FB01F907" lastPageId="17" lastPageNumber="1205" pageId="5" pageNumber="1193">
<subSubSection id="C35B650B2270B05A2565EF19FD4FF9A7" box="[279,662,1620,1643]" pageId="5" pageNumber="1193" type="nomenclature">
<paragraph id="8BFE36802270B05A2565EF19FD4FF9A7" blockId="5.[279,662,1620,1675]" box="[279,662,1620,1643]" pageId="5" pageNumber="1193">
<heading id="D0B681EC2270B05A2565EF19FD4FF9A7" box="[279,662,1620,1643]" centered="true" fontSize="10" level="2" pageId="5" pageNumber="1193" reason="8">
<taxonomicName id="4C414D032270B05A2551EF18FD4FF9A7" authority="(Jaekel, 1894)" baseAuthorityName="Jaekel" baseAuthorityYear="1894" box="[291,662,1620,1643]" class="Elasmobranchii" family="Narcinidae" genus="Titanonarke" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="5" pageNumber="1193" phylum="Chordata" rank="species" species="molini">
<emphasis id="B935EA922270B05A2551EF18FE21F9A7" bold="true" box="[291,504,1620,1643]" italics="true" pageId="5" pageNumber="1193">Titanonarke molini</emphasis>
(
<bibRefCitation id="EFD04B712270B05A2674EF18FD55F9A7" author="Jaekel, O." box="[518,652,1620,1643]" pageId="5" pageNumber="1193" refId="ref23357" refString="Jaekel, O. 1894. Die eoca ¨ nen Selachier vom Monte Bolca: ein Beitrag zur Morphogenie der Wirbelthiere. Springer, Berlin, 177 pp." type="book" year="1894">Jaekel, 1894</bibRefCitation>
)
</taxonomicName>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="C35B650B2270B05A25EBEF38FC4EFCDE" pageId="5" pageNumber="1193" type="description">
<paragraph id="8BFE36802270B05A25EBEF38FDCDF947" blockId="5.[279,662,1620,1675]" box="[409,532,1652,1675]" pageId="5" pageNumber="1193">
(
<figureCitation id="137A2A052270B05A25D0EF38FDD2F947" box="[418,523,1652,1675]" captionStart-0="Figure 2" captionStart-1="Figure 3" captionStart-10="Figure 12" captionStart-11="Figure 13" captionStart-12="Figure 14" captionStart-13="Figure 15" captionStart-14="Figure 16" captionStart-2="Figure 4" captionStart-3="Figure 5" captionStart-4="Figure 6" captionStart-5="Figure 7" captionStart-6="Figure 8" captionStart-7="Figure 9" captionStart-8="Figure 10" captionStart-9="Figure 11" captionStartId-0="6.[159,227,1158,1179]" captionStartId-1="7.[159,227,1005,1026]" captionStartId-10="11.[159,227,1780,1801]" captionStartId-11="12.[159,227,761,782]" captionStartId-12="12.[159,227,1806,1827]" captionStartId-13="13.[159,227,1162,1183]" captionStartId-14="14.[159,227,1045,1066]" captionStartId-2="7.[159,227,1824,1845]" captionStartId-3="8.[159,227,682,703]" captionStartId-4="8.[159,227,1806,1827]" captionStartId-5="9.[159,227,863,884]" captionStartId-6="9.[159,227,1833,1854]" captionStartId-7="10.[159,227,702,723]" captionStartId-8="10.[159,227,1833,1854]" captionStartId-9="11.[159,227,701,722]" captionTargetBox-0="[365,1261,183,1126]" captionTargetBox-1="[365,1261,183,973]" captionTargetBox-10="[166,774,1083,1748]" captionTargetBox-11="[317,1309,183,729]" captionTargetBox-12="[166,774,1108,1775]" captionTargetBox-13="[173,1451,183,1130]" captionTargetBox-14="[173,1453,183,1013]" captionTargetBox-2="[365,1261,1139,1791]" captionTargetBox-3="[365,1255,183,650]" captionTargetBox-4="[365,1252,1347,1774]" captionTargetBox-5="[317,1309,183,831]" captionTargetBox-6="[318,1308,1256,1800]" captionTargetBox-7="[365,1261,183,670]" captionTargetBox-8="[365,1248,1190,1801]" captionTargetBox-9="[365,1257,183,669]" captionTargetId-0="figure-390@6.[365,1261,183,1126]" captionTargetId-1="figure-1@7.[365,1261,183,973]" captionTargetId-10="figure-435@11.[166,774,1083,1748]" captionTargetId-11="figure-291@12.[317,1309,183,729]" captionTargetId-12="figure-371@12.[166,774,1108,1775]" captionTargetId-13="figure-322@13.[173,1453,183,1130]" captionTargetId-14="figure-382@14.[173,1453,183,1013]" captionTargetId-2="figure-28@7.[365,1261,1139,1791]" captionTargetId-3="figure-236@8.[365,1261,183,650]" captionTargetId-4="figure-317@8.[365,1261,1347,1774]" captionTargetId-5="figure-95@9.[317,1309,183,831]" captionTargetId-6="figure-211@9.[317,1309,1255,1801]" captionTargetId-7="figure-180@10.[365,1261,183,670]" captionTargetId-8="figure-248@10.[365,1261,1190,1801]" captionTargetId-9="figure-384@11.[365,1261,183,669]" captionTargetPageId-0="6" captionTargetPageId-1="7" captionTargetPageId-10="11" captionTargetPageId-11="12" captionTargetPageId-12="12" captionTargetPageId-13="13" captionTargetPageId-14="14" captionTargetPageId-2="7" captionTargetPageId-3="8" captionTargetPageId-4="8" captionTargetPageId-5="9" captionTargetPageId-6="9" captionTargetPageId-7="10" captionTargetPageId-8="10" captionTargetPageId-9="11" captionText-0="Figure 2. †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site. A, B, MGP-PD 26275/6, holotype in part and counterpart. Scale bars = 100 mm." captionText-1="Figure 3. †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site. A, MCSNV IG.VR.67290; B, MCSNV IG.91128/9. Scale bars = 100 mm." captionText-10="Figure 12. †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site. Close-up of the distal end of pelvic fins of MCSNV IG.91128, which is supposed to be the unique male individual based on the presence of claspers. Abbreviations: cla, clasper; dfr, first dorsal-fin radials, pelvic radials. Scale bar = 50 mm." captionText-11="Figure 13. †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site. Details of the precaudal tail of A, MCSNV IG. VR.67290 and B, MGP-PD 26276. The arrows mark the position of the two dorsal fins in MCSNV IG.VR.67290 and the second dorsal fin in MGP-PD 26276 already detected and figured by Jaekel (1894). C, Detail of the caudal fin of MCSNV IG.91129. Scale bars: A, B = 50 mm; C = 10 mm." captionText-12="Figure 14. †Titanonarke molini (Jaekel, 1894) from the Eocene of Monte Postale site. A, upper and lower tooth bands in MCSNV IG.VR.67290, with a close-up of some teeth in the area indicated. B, reconstruction. Abbreviations: la, labial cartilage; me, Meckels cartilage; pq, palatoquadrate. Scale bars 5 mm." captionText-13="Figure 15. A, B, †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site; A, detail of the abdominal region in MGP-PD 26275 showing the stomach content; B, reconstruction; note also the embryo lying next to the stomach. C, D, close-up of some of the larger foraminifera of the genus †Alveolina in the stomach of MGP-PD 26275. E, dissected specimen of Torpedo nobiliana in ventral view (ESB tn200707_159) showing the position of the stomach, used to identify the accumulation in MGP-PD 26275 as gut contents. Abbreviations: int, intestine; liv, liver; met, metapterygium; pub, puboischiadic bar; sca, scapulocoracoid; st, stomach, syn synarcual. Scale bars: A, B = 50 mm; C, D = 2 mm." captionText-14="Figure 16. A, B, D, †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site; A, close-up of the abdominal region of MGP-PD 26275 showing the embryo; the anterior region of the body lies on the lower portion of the photo; B, reconstruction; D, detail of the vertebral column of the embryo indicated in B with a dotted rectangle. C, dissected specimen of Potamotrygon tigrina (IUWP 7361) showing the position of the left uterus, just next to the stomach. Abbreviations: int, intestine; liv, liver; lu, left uterus; na, neural arches; ru, right uterus; st, stomach; vc, vertebral centra. Scale bars: A, B = 10 mm; D = 1 mm." captionText-2="Figure 4. Juvenile individuals of †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site. A, MCSNV IG.VR.91359; B, MCSNV IG.135581. Scale bars = 10 mm." captionText-3="Figure 5. †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site. A, MCSNV IG.VR.67290, close-up of the head and hyoid apparatus; B, reconstruction. Abbreviations: ao, antorbital cartilage; bb, basibranchials; bbc, basibranchial copula; cb, ceratobranchials; cc, chondrocranium; eb, epibranchials; hb, hypobranchials; hym, hyomandibula; la, labial cartilages; me, Meckels cartilage; nc, nasal capsule; pq, palatoquadrate; ps, pseudohyoid; ra, rostral appendix; rf, rostral fontanelle; ro, rostral cartilage; sca, scapulocoracoid; ss, suprascapula; syn, synarcual. Scale bars = 50 mm." captionText-4="Figure 6. †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site. A, the juvenile individual MCSNV IG.VR.91359; B, reconstruction. Abbreviations: ao, antorbital cartilage; cb, ceratobranchials; hym, hyomandibula; la, labial cartilages; me, Meckels cartilage; mes, mesopterygium; met, metapterygium; nc, nasal capsule; pel, prepelvic process; pub, puboischiadic bar; pq, palatoquadrate; pro, propterygium; ps, pseudohyoid; rad, pectoral radials; ro, rostral cartilage; sca, scapulocoracoid; ss, suprascapula; syn, synarcual. Scale bars = 50 mm." captionText-5="Figure 7. †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site. A, B, UV images of the holotype MGP-PD 26275/6 showing that it was covered with a pigment reflecting an orange light (see also Supplementary material). The arrows in A indicate a leaf also covered with the pigment, a drop, and some rays that were not distally covered (they are blue/grey, as expected); the arrowhead in A indicates part of the third branch of the antorbital cartilage not covered by pigment; the arrowhead in B indicates the rostral appendix; C, Narcine brasiliensis (TNHC 18512); the arrows indicate the rostral appendices. Scale bars: A, B = 50 mm; C = 10 mm." captionText-6="Figure 8. Visual image of the principal component analysis (PCA) performed on the entire set of log-transformed standardized morphometric and meristic features, showing the separation of the specimens referred to †Titanonarke molini (right side of morphospace) from those referred to †Titanonarke megapterygia sp. nov. (left side). The illustrations lying along the extreme values of PC1 represent the hypothetical reconstruction of the two species based on body proportions." captionText-7="Figure 9. A, synarcual and pectoral girdle of Narcine brasiliensis (AMNH 95343) in dorsal view; B, synarcual and pectoral girdle of †Titanonarke molini (MCSNV IG.VR.91359) in ventral view. The arrowheads indicate the posteriorly directed lateral stays of the synarcual. The arrows indicate the posteriorly directed scapular process of the scapulocoracoid. Abbreviations: cb, ceratocranchials; sca, scapulocoracoid; ss, suprascapula; syn, synarcual. Scale bar = 5 mm." captionText-8="Figure 10. Pectoral fin of †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site. A, MCSNV IG.VR.67290; B, reconstruction; different colours are used to distinguish the propterygial (green), mesopterygial (yellow) and metapterygial (red) radials (colours in the online version). Abbreviations: mes, mesopterygium; met, metapterygium; pro, propterygium; sca, scapulocoracoid. Scale bars = 50 mm." captionText-9="Figure 11. Pelvic fins of †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site. A, MCSNV IG.VR.67290; B, reconstruction. Abbreviations: bas, basipterygia; ilp, iliac process; pel, pelvic processes; pub, puboischiadic bar; rad, pelvic radials. The arrowhead indicates the iliac process. Scale bars = 50 mm." figureDoi-0="http://doi.org/10.5281/zenodo.10883270" figureDoi-1="http://doi.org/10.5281/zenodo.10883273" figureDoi-10="http://doi.org/10.5281/zenodo.10883299" figureDoi-11="http://doi.org/10.5281/zenodo.10883301" figureDoi-12="http://doi.org/10.5281/zenodo.10883305" figureDoi-13="http://doi.org/10.5281/zenodo.10883307" figureDoi-14="http://doi.org/10.5281/zenodo.10883311" figureDoi-2="http://doi.org/10.5281/zenodo.10883277" figureDoi-3="http://doi.org/10.5281/zenodo.10883279" figureDoi-4="http://doi.org/10.5281/zenodo.10883285" figureDoi-5="http://doi.org/10.5281/zenodo.10883287" figureDoi-6="http://doi.org/10.5281/zenodo.10883289" figureDoi-7="http://doi.org/10.5281/zenodo.10883293" figureDoi-8="http://doi.org/10.5281/zenodo.10883295" figureDoi-9="http://doi.org/10.5281/zenodo.10883297" httpUri-0="https://zenodo.org/record/10883270/files/figure.png" httpUri-1="https://zenodo.org/record/10883273/files/figure.png" httpUri-10="https://zenodo.org/record/10883299/files/figure.png" httpUri-11="https://zenodo.org/record/10883301/files/figure.png" httpUri-12="https://zenodo.org/record/10883305/files/figure.png" httpUri-13="https://zenodo.org/record/10883307/files/figure.png" httpUri-14="https://zenodo.org/record/10883311/files/figure.png" httpUri-2="https://zenodo.org/record/10883277/files/figure.png" httpUri-3="https://zenodo.org/record/10883279/files/figure.png" httpUri-4="https://zenodo.org/record/10883285/files/figure.png" httpUri-5="https://zenodo.org/record/10883287/files/figure.png" httpUri-6="https://zenodo.org/record/10883289/files/figure.png" httpUri-7="https://zenodo.org/record/10883293/files/figure.png" httpUri-8="https://zenodo.org/record/10883295/files/figure.png" httpUri-9="https://zenodo.org/record/10883297/files/figure.png" pageId="5" pageNumber="1193">Figs 216</figureCitation>
)
</paragraph>
<paragraph id="8BFE36802270B05A24EDEFF8FD44F907" blockId="5.[159,781,1716,1930]" box="[159,669,1716,1739]" pageId="5" pageNumber="1193">
1860 †
<taxonomicName id="4C414D032270B05A2495EFF8FE35F907" authority="Molin" authorityName="Molin" box="[231,492,1716,1739]" class="Elasmobranchii" family="Narcinidae" genus="Narcine" kingdom="Animalia" order="Torpediniformes" pageId="5" pageNumber="1193" phylum="Chordata" rank="species" species="gigantea">
<emphasis id="B935EA922270B05A2495EFF8FE79F907" box="[231,416,1716,1739]" italics="true" pageId="5" pageNumber="1193">Narcine gigantea</emphasis>
Molin
</taxonomicName>
: 585 [pro parte].
</paragraph>
<paragraph id="8BFE36802270B05A24EDEF9FFCDDF8C6" blockId="5.[159,781,1716,1930]" pageId="5" pageNumber="1193">
1894 †
<taxonomicName id="4C414D032270B05A2498EF9FFE39F926" authority="Jaekel" authorityName="Jaekel" authorityYear="1894" box="[234,480,1747,1770]" class="Elasmobranchii" family="Narcinidae" genus="Narcine" kingdom="Animalia" order="Torpediniformes" pageId="5" pageNumber="1193" phylum="Chordata" rank="species" species="molini">
<emphasis id="B935EA922270B05A2498EF9FFE57F926" box="[234,398,1747,1770]" italics="true" pageId="5" pageNumber="1193">Narcine molini</emphasis>
Jaekel
</taxonomicName>
: 111, pl. 3 [original occurrence of name, photograph and outline reconstruction].
</paragraph>
<paragraph id="8BFE36802270B05A24EDEE5FFD75F8E6" blockId="5.[159,781,1716,1930]" box="[159,684,1811,1834]" pageId="5" pageNumber="1193">
1904 †
<taxonomicName id="4C414D032270B05A2495EE5FFDCBF8E6" authority="Jaekel, 1894" authorityName="Jaekel" authorityYear="1894" box="[231,530,1811,1834]" class="Elasmobranchii" family="Narcinidae" genus="Narcine" kingdom="Animalia" order="Torpediniformes" pageId="5" pageNumber="1193" phylum="Chordata" rank="species" species="molini">
<emphasis id="B935EA922270B05A2495EE5FFE5EF8E6" box="[231,391,1811,1834]" italics="true" pageId="5" pageNumber="1193">Narcine molini</emphasis>
<bibRefCitation id="EFD04B712270B05A25FCEE5FFDCBF8E6" author="Jaekel, O." box="[398,530,1811,1834]" pageId="5" pageNumber="1193" refId="ref23357" refString="Jaekel, O. 1894. Die eoca ¨ nen Selachier vom Monte Bolca: ein Beitrag zur Morphogenie der Wirbelthiere. Springer, Berlin, 177 pp." type="book" year="1894">Jaekel, 1894</bibRefCitation>
</taxonomicName>
; Eastman: 27.
</paragraph>
<paragraph id="8BFE36802270B05A24EDEE7FFD63F886" blockId="5.[159,781,1716,1930]" box="[159,698,1843,1866]" pageId="5" pageNumber="1193">
1905 †
<taxonomicName id="4C414D032270B05A2495EE7FFDCBF886" authority="Jaekel, 1894" authorityName="Jaekel" authorityYear="1894" box="[231,530,1843,1866]" class="Elasmobranchii" family="Narcinidae" genus="Narcine" kingdom="Animalia" order="Torpediniformes" pageId="5" pageNumber="1193" phylum="Chordata" rank="species" species="molini">
<emphasis id="B935EA922270B05A2495EE7FFE5EF886" box="[231,391,1843,1866]" italics="true" pageId="5" pageNumber="1193">Narcine molini</emphasis>
<bibRefCitation id="EFD04B712270B05A25FCEE7FFDCBF886" author="Jaekel, O." box="[398,530,1843,1866]" pageId="5" pageNumber="1193" refId="ref23357" refString="Jaekel, O. 1894. Die eoca ¨ nen Selachier vom Monte Bolca: ein Beitrag zur Morphogenie der Wirbelthiere. Springer, Berlin, 177 pp." type="book" year="1894">Jaekel, 1894</bibRefCitation>
</taxonomicName>
; Eastman: 351.
</paragraph>
<paragraph id="8BFE36802270B05A24EDEE1FFE54F846" blockId="5.[159,781,1716,1930]" pageId="5" pageNumber="1193">
1979 †
<taxonomicName id="4C414D032270B05A2480EE1FFDE6F8A6" authority="Jaekel, 1894" authorityName="Jaekel" authorityYear="1894" box="[242,575,1875,1898]" class="Elasmobranchii" family="Narcinidae" genus="Narcine" kingdom="Animalia" order="Torpediniformes" pageId="5" pageNumber="1193" phylum="Chordata" rank="species" species="molini">
<emphasis id="B935EA922270B05A2480EE1FFE44F8A6" box="[242,413,1875,1898]" italics="true" pageId="5" pageNumber="1193">Narcine molini</emphasis>
<bibRefCitation id="EFD04B712270B05A25DDEE1FFDE6F8A6" author="Jaekel, O." box="[431,575,1875,1898]" pageId="5" pageNumber="1193" refId="ref23357" refString="Jaekel, O. 1894. Die eoca ¨ nen Selachier vom Monte Bolca: ein Beitrag zur Morphogenie der Wirbelthiere. Springer, Berlin, 177 pp." type="book" year="1894">Jaekel, 1894</bibRefCitation>
</taxonomicName>
; Stanghellini: 38 [misspelt
<emphasis id="B935EA922270B05A2559EE3FFEA7F846" box="[299,382,1907,1930]" italics="true" pageId="5" pageNumber="1193">moloni</emphasis>
].
</paragraph>
<paragraph id="8BFE36802270B05A273FE9F1FAE0FF18" blockId="5.[845,1467,189,786]" box="[845,1337,189,212]" pageId="5" pageNumber="1193">
1980 †
<taxonomicName id="4C414D032270B05A27E6E9F1FB19FF18" authority="Jaekel, 1894" authorityName="Jaekel" authorityYear="1894" box="[916,1216,189,212]" class="Elasmobranchii" family="Narcinidae" genus="Narcine" kingdom="Animalia" order="Torpediniformes" pageId="5" pageNumber="1193" phylum="Chordata" rank="species" species="molini">
<emphasis id="B935EA922270B05A27E6E9F1FBEDFF18" box="[916,1076,189,212]" italics="true" pageId="5" pageNumber="1193">Narcine molini</emphasis>
<bibRefCitation id="EFD04B712270B05A2049E9F1FB19FF18" author="Jaekel, O." box="[1083,1216,189,212]" pageId="5" pageNumber="1193" refId="ref23357" refString="Jaekel, O. 1894. Die eoca ¨ nen Selachier vom Monte Bolca: ein Beitrag zur Morphogenie der Wirbelthiere. Springer, Berlin, 177 pp." type="book" year="1894">Jaekel, 1894</bibRefCitation>
</taxonomicName>
; Blot: 344.
</paragraph>
<paragraph id="8BFE36802270B05A273FE991FC09FED8" blockId="5.[845,1467,189,786]" pageId="5" pageNumber="1193">
1987 †
<taxonomicName id="4C414D032270B05A27D0E991FB2FFF38" authority="Jaekel, 1894" authorityName="Jaekel" authorityYear="1894" box="[930,1270,221,244]" class="Elasmobranchii" family="Narcinidae" genus="Narcine" kingdom="Animalia" order="Torpediniformes" pageId="5" pageNumber="1193" phylum="Chordata" rank="species" species="molini">
<emphasis id="B935EA922270B05A27D0E991FB96FF38" box="[930,1103,221,244]" italics="true" pageId="5" pageNumber="1193">Narcine molini</emphasis>
<bibRefCitation id="EFD04B712270B05A2011E991FB2FFF38" author="Jaekel, O." box="[1123,1270,221,244]" pageId="5" pageNumber="1193" refId="ref23357" refString="Jaekel, O. 1894. Die eoca ¨ nen Selachier vom Monte Bolca: ein Beitrag zur Morphogenie der Wirbelthiere. Springer, Berlin, 177 pp." type="book" year="1894">Jaekel, 1894</bibRefCitation>
</taxonomicName>
; Cappetta: 161, fig.
<specimenCount id="9D47FD092270B05A27E2E9B1FC09FED8" box="[912,976,253,276]" count="138" pageId="5" pageNumber="1193" type="generic">138L.</specimenCount>
</paragraph>
<paragraph id="8BFE36802270B05A273FE851FA84FEF8" blockId="5.[845,1467,189,786]" box="[845,1373,285,308]" pageId="5" pageNumber="1193">
1988 †
<taxonomicName id="4C414D032270B05A27E6E851FB19FEF8" authority="Jaekel, 1894" authorityName="Jaekel" authorityYear="1894" box="[916,1216,285,308]" class="Elasmobranchii" family="Narcinidae" genus="Narcine" kingdom="Animalia" order="Torpediniformes" pageId="5" pageNumber="1193" phylum="Chordata" rank="species" species="molini">
<emphasis id="B935EA922270B05A27E6E851FBEDFEF8" box="[916,1076,285,308]" italics="true" pageId="5" pageNumber="1193">Narcine molini</emphasis>
<bibRefCitation id="EFD04B712270B05A2049E851FB19FEF8" author="Jaekel, O." box="[1083,1216,285,308]" pageId="5" pageNumber="1193" refId="ref23357" refString="Jaekel, O. 1894. Die eoca ¨ nen Selachier vom Monte Bolca: ein Beitrag zur Morphogenie der Wirbelthiere. Springer, Berlin, 177 pp." type="book" year="1894">Jaekel, 1894</bibRefCitation>
</taxonomicName>
; Cappetta: 29.
</paragraph>
<paragraph id="8BFE36802270B05A273FE871FA5EFE98" blockId="5.[845,1467,189,786]" box="[845,1415,317,340]" pageId="5" pageNumber="1193">
1991 †
<taxonomicName id="4C414D032270B05A27E6E871FB19FE98" authority="Jaekel, 1894" authorityName="Jaekel" authorityYear="1894" box="[916,1216,317,340]" class="Elasmobranchii" family="Narcinidae" genus="Narcine" kingdom="Animalia" order="Torpediniformes" pageId="5" pageNumber="1193" phylum="Chordata" rank="species" species="molini">
<emphasis id="B935EA922270B05A27E6E871FBEDFE98" box="[916,1076,317,340]" italics="true" pageId="5" pageNumber="1193">Narcine molini</emphasis>
<bibRefCitation id="EFD04B712270B05A2049E871FB19FE98" author="Jaekel, O." box="[1083,1216,317,340]" pageId="5" pageNumber="1193" refId="ref23357" refString="Jaekel, O. 1894. Die eoca ¨ nen Selachier vom Monte Bolca: ein Beitrag zur Morphogenie der Wirbelthiere. Springer, Berlin, 177 pp." type="book" year="1894">Jaekel, 1894</bibRefCitation>
</taxonomicName>
; Frickhinger: 211.
</paragraph>
<paragraph id="8BFE36802270B05A273FE811FA7FFEB8" blockId="5.[845,1467,189,786]" box="[845,1446,349,372]" pageId="5" pageNumber="1193">
1993 †
<taxonomicName id="4C414D032270B05A27E6E811FB19FEB8" authority="Jaekel, 1894" authorityName="Jaekel" authorityYear="1894" box="[916,1216,349,372]" class="Elasmobranchii" family="Narcinidae" genus="Narcine" kingdom="Animalia" order="Torpediniformes" pageId="5" pageNumber="1193" phylum="Chordata" rank="species" species="molini">
<emphasis id="B935EA922270B05A27E6E811FBEDFEB8" box="[916,1076,349,372]" italics="true" pageId="5" pageNumber="1193">Narcine molini</emphasis>
<bibRefCitation id="EFD04B712270B05A2049E811FB19FEB8" author="Jaekel, O." box="[1083,1216,349,372]" pageId="5" pageNumber="1193" refId="ref23357" refString="Jaekel, O. 1894. Die eoca ¨ nen Selachier vom Monte Bolca: ein Beitrag zur Morphogenie der Wirbelthiere. Springer, Berlin, 177 pp." type="book" year="1894">Jaekel, 1894</bibRefCitation>
</taxonomicName>
; Cappetta
<emphasis id="B935EA922270B05A2141E811FAB0FEB8" box="[1331,1385,349,372]" italics="true" pageId="5" pageNumber="1193">et al.</emphasis>
: 604.
</paragraph>
<paragraph id="8BFE36802270B05A273FE831FC21FE78" blockId="5.[845,1467,189,786]" pageId="5" pageNumber="1193">
1999 †‘
<taxonomicName id="4C414D032270B05A27D5E831FB23FE58" authority="Jaekel, 1894" authorityName="Jaekel" authorityYear="1894" box="[935,1274,381,404]" class="Elasmobranchii" family="Narcinidae" genus="Narcine" kingdom="Animalia" order="Torpediniformes" pageId="5" pageNumber="1193" phylum="Chordata" rank="species" species="molini">
<emphasis id="B935EA922270B05A27D5E831FC24FE58" box="[935,1021,381,404]" italics="true" pageId="5" pageNumber="1193">Narcine</emphasis>
<emphasis id="B935EA922270B05A2064E831FB83FE58" box="[1046,1114,381,404]" italics="true" pageId="5" pageNumber="1193">molini</emphasis>
<bibRefCitation id="EFD04B712270B05A2019E831FB23FE58" author="Jaekel, O." box="[1131,1274,381,404]" pageId="5" pageNumber="1193" refId="ref23357" refString="Jaekel, O. 1894. Die eoca ¨ nen Selachier vom Monte Bolca: ein Beitrag zur Morphogenie der Wirbelthiere. Springer, Berlin, 177 pp." type="book" year="1894">Jaekel, 1894</bibRefCitation>
</taxonomicName>
; Carvalho: 283, figs 107111.
</paragraph>
<paragraph id="8BFE36802270B05A273FE8F0FBD3FE3F" blockId="5.[845,1467,189,786]" pageId="5" pageNumber="1193">
2010 †
<taxonomicName id="4C414D032270B05A27E5E8F0FAD2FE1F" authority="(Jaekel, 1894)" baseAuthorityName="Jaekel" baseAuthorityYear="1894" box="[919,1291,444,467]" class="Elasmobranchii" family="Narcinidae" genus="Titanonarke" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="5" pageNumber="1193" phylum="Chordata" rank="species" species="molini">
<emphasis id="B935EA922270B05A27E5E8F0FBBFFE1F" box="[919,1126,444,467]" italics="true" pageId="5" pageNumber="1193">Titanonarke molini</emphasis>
(
<bibRefCitation id="EFD04B712270B05A2005E8F0FAD9FE1F" author="Jaekel, O." box="[1143,1280,444,467]" pageId="5" pageNumber="1193" refId="ref23357" refString="Jaekel, O. 1894. Die eoca ¨ nen Selachier vom Monte Bolca: ein Beitrag zur Morphogenie der Wirbelthiere. Springer, Berlin, 177 pp." type="book" year="1894">Jaekel, 1894</bibRefCitation>
)
</taxonomicName>
; Carvalho: 185, figs 2, 5A, 6, 7.
</paragraph>
<paragraph id="8BFE36802270B05A273FE8B0FC15FDFF" blockId="5.[845,1467,189,786]" pageId="5" pageNumber="1193">
2012b †
<taxonomicName id="4C414D032270B05A27D9E8B0FAEBFDDF" authority="(Jaekel, 1894)" baseAuthorityName="Jaekel" baseAuthorityYear="1894" box="[939,1330,508,531]" class="Elasmobranchii" family="Narcinidae" genus="Titanonarke" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="5" pageNumber="1193" phylum="Chordata" rank="species" species="molini">
<emphasis id="B935EA922270B05A27D9E8B0FB59FDDF" box="[939,1152,508,531]" italics="true" pageId="5" pageNumber="1193">Titanonarke molini</emphasis>
(
<bibRefCitation id="EFD04B712270B05A20E5E8B0FAFEFDDF" author="Jaekel, O." box="[1175,1319,508,531]" pageId="5" pageNumber="1193" refId="ref23357" refString="Jaekel, O. 1894. Die eoca ¨ nen Selachier vom Monte Bolca: ein Beitrag zur Morphogenie der Wirbelthiere. Springer, Berlin, 177 pp." type="book" year="1894">Jaekel, 1894</bibRefCitation>
)
</taxonomicName>
; Aschliman
<emphasis id="B935EA922270B05A2715EB50FC44FDFF" box="[871,925,540,563]" italics="true" pageId="5" pageNumber="1193">et al.</emphasis>
: 33.
</paragraph>
<paragraph id="8BFE36802270B05A273FEB70FC09FDBF" blockId="5.[845,1467,189,786]" pageId="5" pageNumber="1193">
2012 †
<taxonomicName id="4C414D032270B05A27D0EB70FB2FFD9F" authority="Jaekel, 1894" authorityName="Jaekel" authorityYear="1894" box="[930,1270,572,595]" class="Elasmobranchii" family="Narcinidae" genus="Narcine" kingdom="Animalia" order="Torpediniformes" pageId="5" pageNumber="1193" phylum="Chordata" rank="species" species="molini">
<emphasis id="B935EA922270B05A27D0EB70FB96FD9F" box="[930,1103,572,595]" italics="true" pageId="5" pageNumber="1193">Narcine molini</emphasis>
<bibRefCitation id="EFD04B712270B05A2011EB70FB2FFD9F" author="Jaekel, O." box="[1123,1270,572,595]" pageId="5" pageNumber="1193" refId="ref23357" refString="Jaekel, O. 1894. Die eoca ¨ nen Selachier vom Monte Bolca: ein Beitrag zur Morphogenie der Wirbelthiere. Springer, Berlin, 177 pp." type="book" year="1894">Jaekel, 1894</bibRefCitation>
</taxonomicName>
; Cappetta: 410, fig.
<specimenCount id="9D47FD092270B05A27E2EB10FC09FDBF" box="[912,976,604,627]" count="401" pageId="5" pageNumber="1193" type="generic">401L.</specimenCount>
</paragraph>
<paragraph id="8BFE36802270B05A273FEB30FC50FD7F" blockId="5.[845,1467,189,786]" pageId="5" pageNumber="1193">
2014 †
<taxonomicName id="4C414D032270B05A27E6EB30FAD9FD5F" authority="(Jaekel, 1894)" baseAuthorityName="Jaekel" baseAuthorityYear="1894" box="[916,1280,636,659]" class="Elasmobranchii" family="Narcinidae" genus="Titanonarke" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="5" pageNumber="1193" phylum="Chordata" rank="species" species="molini">
<emphasis id="B935EA922270B05A27E6EB30FBB8FD5F" box="[916,1121,636,659]" italics="true" pageId="5" pageNumber="1193">Titanonarke molini</emphasis>
(
<bibRefCitation id="EFD04B712270B05A201CEB30FB2CFD5F" author="Jaekel, O." box="[1134,1269,636,659]" pageId="5" pageNumber="1193" refId="ref23357" refString="Jaekel, O. 1894. Die eoca ¨ nen Selachier vom Monte Bolca: ein Beitrag zur Morphogenie der Wirbelthiere. Springer, Berlin, 177 pp." type="book" year="1894">Jaekel, 1894</bibRefCitation>
)
</taxonomicName>
; Carnevale
<emphasis id="B935EA922270B05A210CEB30FA6AFD5F" box="[1406,1459,636,659]" italics="true" pageId="5" pageNumber="1193">et al.</emphasis>
: 41.
</paragraph>
<paragraph id="8BFE36802270B05A273FEBF7FA5DFD1E" blockId="5.[845,1467,189,786]" box="[845,1412,699,722]" pageId="5" pageNumber="1193">
2014 †
<taxonomicName id="4C414D032270B05A27E6EBF7FAD9FD1E" authority="(Jaekel, 1894)" baseAuthorityName="Jaekel" baseAuthorityYear="1894" box="[916,1280,699,722]" class="Elasmobranchii" family="Narcinidae" genus="Titanonarke" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="5" pageNumber="1193" phylum="Chordata" rank="species" species="molini">
<emphasis id="B935EA922270B05A27E6EBF7FBB8FD1E" box="[916,1121,699,722]" italics="true" pageId="5" pageNumber="1193">Titanonarke molini</emphasis>
(
<bibRefCitation id="EFD04B712270B05A201DEBF7FB2CFD1E" author="Jaekel, O." box="[1135,1269,699,722]" pageId="5" pageNumber="1193" refId="ref23357" refString="Jaekel, O. 1894. Die eoca ¨ nen Selachier vom Monte Bolca: ein Beitrag zur Morphogenie der Wirbelthiere. Springer, Berlin, 177 pp." type="book" year="1894">Jaekel, 1894</bibRefCitation>
)
</taxonomicName>
; Claeson: 4.
</paragraph>
<paragraph id="8BFE36802270B05A273FEB97FC4EFCDE" blockId="5.[845,1467,189,786]" pageId="5" pageNumber="1193">
2016c †
<taxonomicName id="4C414D032270B05A27D2EB97FAD2FD3E" authority="(Jaekel, 1894)" baseAuthorityName="Jaekel" baseAuthorityYear="1894" box="[928,1291,731,754]" class="Elasmobranchii" family="Narcinidae" genus="Titanonarke" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="5" pageNumber="1193" phylum="Chordata" rank="species" species="molini">
<emphasis id="B935EA922270B05A27D2EB97FBB5FD3E" box="[928,1132,731,754]" italics="true" pageId="5" pageNumber="1193">Titanonarke molini</emphasis>
(
<bibRefCitation id="EFD04B712270B05A2008EB97FAD9FD3E" author="Jaekel, O." box="[1146,1280,731,754]" pageId="5" pageNumber="1193" refId="ref23357" refString="Jaekel, O. 1894. Die eoca ¨ nen Selachier vom Monte Bolca: ein Beitrag zur Morphogenie der Wirbelthiere. Springer, Berlin, 177 pp." type="book" year="1894">Jaekel, 1894</bibRefCitation>
)
</taxonomicName>
; Marram̀a
<emphasis id="B935EA922270B05A210CEB97FA6AFD3E" box="[1406,1459,731,754]" italics="true" pageId="5" pageNumber="1193">et al.</emphasis>
: 232.
</paragraph>
</subSubSection>
<subSubSection id="C35B650B2270B05A273FEA62FB9BFAE6" pageId="5" pageNumber="1193" type="materials_examined">
<paragraph id="8BFE36802270B05A273FEA62FA40FCA9" blockId="5.[845,1466,814,869]" pageId="5" pageNumber="1193">
<materialsCitation id="3B293CDD2270B05A273FEA62FA40FCA9" pageId="5" pageNumber="1193" specimenCode="MGP-PD 26275" specimenCount="1" typeStatus="holotype">
<emphasis id="B935EA922270B05A273FEA62FC62FC89" bold="true" box="[845,955,814,837]" pageId="5" pageNumber="1193">
<typeStatus id="54FA88222270B05A273FEA62FC6EFC89" box="[845,951,814,837]" pageId="5" pageNumber="1193" type="holotype">Holotype</typeStatus>
.
</emphasis>
<specimenCode id="DBE79EFB2270B05A27B8EA62FBA0FC89" box="[970,1145,814,837]" collectionCode="MGP-PD" pageId="5" pageNumber="1193">MGP-PD 26275</specimenCode>
/6, nearly complete articulated skeleton in part and counterpart (
<figureCitation id="137A2A052270B05A20DFEA02FB32FCA9" box="[1197,1259,846,869]" captionStart="Figure 2" captionStartId="6.[159,227,1158,1179]" captionTargetBox="[365,1261,183,1126]" captionTargetId="figure-390@6.[365,1261,183,1126]" captionTargetPageId="6" captionText="Figure 2. †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site. A, B, MGP-PD 26275/6, holotype in part and counterpart. Scale bars = 100 mm." figureDoi="http://doi.org/10.5281/zenodo.10883270" httpUri="https://zenodo.org/record/10883270/files/figure.png" pageId="5" pageNumber="1193">Fig. 2</figureCitation>
), 816.8 mm SL.
</materialsCitation>
</paragraph>
<paragraph id="8BFE36802270B05A273FEACDFC6CFB5B" blockId="5.[845,1467,897,1175]" pageId="5" pageNumber="1193">
<emphasis id="B935EA922270B05A273FEACDFBFAFC54" bold="true" box="[845,1059,897,920]" pageId="5" pageNumber="1193">Referred material.</emphasis>
<collectionCode id="ED50AE452270B05A2043EACDFB56FC54" box="[1073,1167,897,920]" pageId="5" pageNumber="1193">MCSNV</collectionCode>
<collectionCode id="ED50AE452270B05A20EAEACDFB6FFC54" box="[1176,1206,897,920]" country="China" name="Institute of Geology" pageId="5" pageNumber="1193">IG</collectionCode>
.VR.67290, nearly complete articulated skeleton in a single slab, 750.2 mm SL (
<figureCitation id="137A2A052270B05A2724EA8DFC72FC14" box="[854,939,961,984]" captionStart="Figure 3" captionStartId="7.[159,227,1005,1026]" captionTargetBox="[365,1261,183,973]" captionTargetId="figure-1@7.[365,1261,183,973]" captionTargetPageId="7" captionText="Figure 3. †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site. A, MCSNV IG.VR.67290; B, MCSNV IG.91128/9. Scale bars = 100 mm." figureDoi="http://doi.org/10.5281/zenodo.10883273" httpUri="https://zenodo.org/record/10883273/files/figure.png" pageId="5" pageNumber="1193">Fig. 3A</figureCitation>
);
<collectionCode id="ED50AE452270B05A27CCEA8DFBC5FC14" box="[958,1052,961,984]" pageId="5" pageNumber="1193">MCSNV</collectionCode>
<collectionCode id="ED50AE452270B05A2056EA8DFB9BFC14" box="[1060,1090,961,984]" country="China" name="Institute of Geology" pageId="5" pageNumber="1193">IG</collectionCode>
.91128/9, partially complete articulated specimen in part and counterpart, 850.4 mm SL (
<figureCitation id="137A2A052270B05A2724ED4DFC71FBD4" box="[854,936,1025,1048]" captionStart="Figure 3" captionStartId="7.[159,227,1005,1026]" captionTargetBox="[365,1261,183,973]" captionTargetId="figure-1@7.[365,1261,183,973]" captionTargetPageId="7" captionText="Figure 3. †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site. A, MCSNV IG.VR.67290; B, MCSNV IG.91128/9. Scale bars = 100 mm." figureDoi="http://doi.org/10.5281/zenodo.10883273" httpUri="https://zenodo.org/record/10883273/files/figure.png" pageId="5" pageNumber="1193">Fig. 3B</figureCitation>
);
<collectionCode id="ED50AE452270B05A27CFED4DFBC2FBD4" box="[957,1051,1025,1048]" pageId="5" pageNumber="1193">MCSNV</collectionCode>
<collectionCode id="ED50AE452270B05A2051ED4DFB98FBD4" box="[1059,1089,1025,1048]" country="China" name="Institute of Geology" pageId="5" pageNumber="1193">IG</collectionCode>
.VR.91359, nearly complete articulated skeleton in a single slab, 99.0 mm SL (
<figureCitation id="137A2A052270B05A2126ED6DFA77FBF4" box="[1364,1454,1057,1080]" captionStart="Figure 4" captionStartId="7.[159,227,1824,1845]" captionTargetBox="[365,1261,1139,1791]" captionTargetId="figure-28@7.[365,1261,1139,1791]" captionTargetPageId="7" captionText="Figure 4. Juvenile individuals of †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site. A, MCSNV IG.VR.91359; B, MCSNV IG.135581. Scale bars = 10 mm." figureDoi="http://doi.org/10.5281/zenodo.10883277" httpUri="https://zenodo.org/record/10883277/files/figure.png" pageId="5" pageNumber="1193">Fig. 4A</figureCitation>
);
<collectionCode id="ED50AE452270B05A273FED0CFC72FB9B" box="[845,939,1088,1111]" pageId="5" pageNumber="1193">MCSNV</collectionCode>
<collectionCode id="ED50AE452270B05A27C0ED0CFC08FB9B" box="[946,977,1088,1111]" country="China" name="Institute of Geology" pageId="5" pageNumber="1193">IG</collectionCode>
.135581, incomplete articulated specimen in a single slab, lacking part of the cranial and caudal regions (
<figureCitation id="137A2A052270B05A2724EDCCFC71FB5B" box="[854,936,1152,1175]" captionStart="Figure 4" captionStartId="7.[159,227,1824,1845]" captionTargetBox="[365,1261,1139,1791]" captionTargetId="figure-28@7.[365,1261,1139,1791]" captionTargetPageId="7" captionText="Figure 4. Juvenile individuals of †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site. A, MCSNV IG.VR.91359; B, MCSNV IG.135581. Scale bars = 10 mm." figureDoi="http://doi.org/10.5281/zenodo.10883277" httpUri="https://zenodo.org/record/10883277/files/figure.png" pageId="5" pageNumber="1193">Fig. 4B</figureCitation>
).
</paragraph>
<paragraph id="8BFE36802270B05A273FEDFFFB9BFAE6" blockId="5.[845,1466,1203,1322]" pageId="5" pageNumber="1193">
<emphasis id="B935EA922270B05A273FEDFFFB50FB06" bold="true" box="[845,1161,1203,1226]" pageId="5" pageNumber="1193">Type locality and horizon.</emphasis>
Monte Postale site, Bolca Konservat-Lagerst¨atte,
<collectingCountry id="F35676102270B05A203BED9FFBA4FB26" box="[1097,1149,1235,1258]" name="Italy" pageId="5" pageNumber="1193">Italy</collectingCountry>
; early Eocene, late Ypresian, middle Cuisian, SBZ 11,
<taxonomicName id="4C414D032270B05A200AEDBFFA5BFAC6" authority="Zone" authorityName="Zone" box="[1144,1410,1267,1290]" class="Magnoliopsida" family="Loranthaceae" genus="Alveolina" kingdom="Plantae" order="Santalales" pageId="5" pageNumber="1193" phylum="Tracheophyta" rank="species" species="dainelli">
<emphasis id="B935EA922270B05A200AEDBFFAE7FAC6" box="[1144,1342,1267,1290]" italics="true" pageId="5" pageNumber="1193">Alveolina dainelli</emphasis>
Zone
</taxonomicName>
(see Papazzoni
<emphasis id="B935EA922270B05A27B3EC5FFC29FAE6" box="[961,1008,1299,1322]" italics="true" pageId="5" pageNumber="1193">et al</emphasis>
. 2017).
</paragraph>
</subSubSection>
<subSubSection id="C35B650B2270B05A273FEC0AFC15F937" pageId="5" pageNumber="1193" type="diagnosis">
<paragraph id="8BFE36802270B05A273FEC0AFC15F937" blockId="5.[845,1467,1350,1787]" pageId="5" pageNumber="1193">
<emphasis id="B935EA922270B05A273FEC0AFC18FA91" bold="true" box="[845,961,1350,1373]" pageId="5" pageNumber="1193">Diagnosis.</emphasis>
Species of †
<taxonomicName id="4C414D032270B05A2014EC0AFB35FA91" authorityName="Carvalho" authorityYear="2010" box="[1126,1260,1350,1373]" class="Elasmobranchii" family="Narcinidae" genus="Titanonarke" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="5" pageNumber="1193" phylum="Chordata" rank="genus">
<emphasis id="B935EA922270B05A2014EC0AFB35FA91" box="[1126,1260,1350,1373]" italics="true" pageId="5" pageNumber="1193">Titanonarke</emphasis>
</taxonomicName>
characterized by the following combination of characters: subcircular disc length
<emphasis id="B935EA922270B05A27BEECCAFC01FA51" box="[972,984,1414,1437]" italics="true" pageId="5" pageNumber="1193">c</emphasis>
. 44% SL and disc width
<emphasis id="B935EA922270B05A209FECCAFB20FA51" box="[1261,1273,1414,1437]" italics="true" pageId="5" pageNumber="1193">c</emphasis>
. 50% SL; greatly elongated precaudal tail of
<emphasis id="B935EA922270B05A20F4ECE9FB4BFA70" box="[1158,1170,1445,1468]" italics="true" pageId="5" pageNumber="1193">c</emphasis>
. 60% SL; 153155 vertebrae (2730 trunk; 100115 precaudal, 2325 caudal); 1516 tooth rows in upper jaw and 1213 tooth rows in the lower jaw;
<emphasis id="B935EA922270B05A278CEF49FBD3F9D0" box="[1022,1034,1541,1564]" italics="true" pageId="5" pageNumber="1193">c</emphasis>
. 40 pectoral radials (1216 propterygial, 912 mesopterygial, and 1011 metapterygial); single-lobed pelvic fin with 2124 basipterygial radials; first dorsal fin with 79 radials, and second dorsal fin with 67 radials; caudal fin with about 42 radials (20 dorsal and 22 ventral); width of pelvic fins about 50% of disc width; anterior pelvic fin margin length
<emphasis id="B935EA922270B05A213EEF88FA81F917" box="[1356,1368,1732,1755]" italics="true" pageId="5" pageNumber="1193">c</emphasis>
. 24% of disc length.
</paragraph>
</subSubSection>
<subSubSection id="C35B650B2270B04E273FEE5BFB01F907" lastPageId="17" lastPageNumber="1205" pageId="5" pageNumber="1193" type="description">
<paragraph id="8BFE36802270B059273FEE5BFE26F842" blockId="5.[845,1467,1815,1934]" lastBlockId="6.[159,782,1305,1934]" lastPageId="6" lastPageNumber="1194" pageId="5" pageNumber="1193">
<emphasis id="B935EA922270B05A273FEE5BFC0EF8E2" bold="true" box="[845,983,1815,1838]" pageId="5" pageNumber="1193">Description.</emphasis>
Specimens examined comprise different ontogenetic stages and range from 99 to 850.4 mm SL, with the largest specimen reaching
<quantity id="4CB99B652270B05A20BAEE1BFAF9F8A2" box="[1224,1312,1879,1902]" metricMagnitude="-1" metricUnit="m" metricValue="9.25" pageId="5" pageNumber="1193" unit="mm" value="925.0">925 mm</quantity>
TL. Measurements and counts for †
<taxonomicName id="4C414D032270B05A204FEE3BFAD3F842" baseAuthorityName="Jaekel" baseAuthorityYear="1894" box="[1085,1290,1911,1934]" class="Elasmobranchii" family="Narcinidae" genus="Titanonarke" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="5" pageNumber="1193" phylum="Chordata" rank="species" species="molini">
<emphasis id="B935EA922270B05A204FEE3BFAD3F842" box="[1085,1290,1911,1934]" italics="true" pageId="5" pageNumber="1193">Titanonarke molini</emphasis>
</taxonomicName>
are summarized in
<tableCitation id="C6C3033B2273B05924CFEC55FEC5FAFC" box="[189,284,1305,1328]" captionStart="Table 1" captionStartId="15.[159,217,186,207]" captionTargetPageId="15" captionText="Table 1. Morphometric data for all examined specimens of †Titanonarke Carvalho, 2010 from the Eocene Monte Postale site, Bolca Lagerst¨atte. Abbreviations (see the morphological scheme in the Supplemental material): AOW, antorbital cartilage width; CFL, caudal fin length; CFD, caudal fin depth; CLO, clasper length; CPD, caudal peduncle depth; D1B, first dorsal fin base length; D2B, second dorsal fin base length; DCS, dorsal caudal space; DL, disc length; DW, disc width; HL, head length; IDS, interdorsal space; MOW, mouth width; P2A, pelvic fin anterior margin length; P2B, pelvic fin base length; P2S, pelvic fin span; PCS, space from pelvic fin insertion to the caudal fin origin; PD1, predorsal distance up to the first dorsal fin; PD2, predorsal distance up to the second dorsal fin; PDI, pelvic fin insertion to first dorsal fin origin; PDO, pelvic fin origin to first dorsal fin origin; PGW, snout tip to the level of the greatest disc width; PIW, body width at pectoral fin insertions; POR, preoral length; PP2, prepelvic length; SL, standard length; TAL, tail length; TBW, tail base at pelvic fin origin; TL, total length." pageId="6" pageNumber="1194">Tables 1</tableCitation>
and
<tableCitation id="C6C3033B2273B0592527EC55FEBBFAFC" box="[341,354,1305,1328]" captionStart="Table 2" captionStartId="16.[159,217,186,207]" captionTargetPageId="16" captionText="Table 2. Summary of selected meristic features used to discriminate fossil and living genera of the family Narcinidae. Includes new data from the examined material and data from Fechhelm &amp; McEachran (1984), Rincon (1997), Carvalho et al. (1999, 2002a, b, 2003), Carvalho (2001, 2008), Rincon et al. (2001), Carvalho &amp; Śeret (2002), Carvalho &amp; Randall (2003), Menni et al. (2008), Carvalho &amp; White (2016), Last et al. (2016), Froese &amp; Pauly (2017)." pageId="6" pageNumber="1194">2</tableCitation>
. Overall, the body is large and dorsoventrally compressed (
<figureCitation id="137A2A052273B05925EDEC75FE3FFA9C" box="[415,486,1337,1360]" captionStart="Figure 2" captionStartId="6.[159,227,1158,1179]" captionTargetBox="[365,1261,183,1126]" captionTargetId="figure-390@6.[365,1261,183,1126]" captionTargetPageId="6" captionText="Figure 2. †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site. A, B, MGP-PD 26275/6, holotype in part and counterpart. Scale bars = 100 mm." figureDoi="http://doi.org/10.5281/zenodo.10883270" httpUri="https://zenodo.org/record/10883270/files/figure.png" pageId="6" pageNumber="1194">Figs 2</figureCitation>
,
<figureCitation id="137A2A052273B059258AEC75FDDCFA9C" box="[504,517,1337,1360]" captionStart="Figure 3" captionStartId="7.[159,227,1005,1026]" captionTargetBox="[365,1261,183,973]" captionTargetId="figure-1@7.[365,1261,183,973]" captionTargetPageId="7" captionText="Figure 3. †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site. A, MCSNV IG.VR.67290; B, MCSNV IG.91128/9. Scale bars = 100 mm." figureDoi="http://doi.org/10.5281/zenodo.10883273" httpUri="https://zenodo.org/record/10883273/files/figure.png" pageId="6" pageNumber="1194">3</figureCitation>
). The disc is subcircular, slightly ovoid in outline, generally wider than long, and barely overlapping the pelvic fin origin. The largest width of the disc is slightly posterior to its midlength and its edges are continuously curved. The length and width of the disc are about 44% and 50% SL, respectively. The head is approximately 25% of SL and the preoral length is about 9% SL. The precaudal tail is elongated (about 60% SL). †
<taxonomicName id="4C414D032273B05925AFEF74FD69F983" baseAuthorityName="Jaekel" baseAuthorityYear="1894" box="[477,688,1592,1615]" class="Elasmobranchii" family="Narcinidae" genus="Titanonarke" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="6" pageNumber="1194" phylum="Chordata" rank="species" species="molini">
<emphasis id="B935EA922273B05925AFEF74FD69F983" box="[477,688,1592,1615]" italics="true" pageId="6" pageNumber="1194">Titanonarke molini</emphasis>
</taxonomicName>
has two dorsal fins; the predorsal distance at the first dorsal fin is about 62% SL, whereas the predorsal distance at the second dorsal fin is about 78% SL; the interdorsal distance is about 7% SL. The body is totally naked, lacking denticles and thorns. The skeleton is highly calcified and most of the skeletal elements, including parts of the rostral cartilage, jaws, hyomandibulae, antorbital cartilages, synarcual, and pectoral and pelvic girdles, show the typical prismatic calcification of elasmobranch fishes (
<bibRefCitation id="EFD04B712273B059249EEE3BFE36F842" author="Dean, M. N. &amp; Summers, A. P." box="[236,495,1911,1934]" pageId="6" pageNumber="1194" pagination="164 - 168" refId="ref21150" refString="Dean, M. N. &amp; Summers, A. P. 2006. Mineralized cartilage in the skeleton of chondrichthyan fishes. Zoology, 109, 164 - 168." type="journal article" year="2006">Dean &amp; Summers 2006</bibRefCitation>
).
</paragraph>
<caption id="DF3E66082273B05924EDEDCAFE19FB7A" ID-DOI="http://doi.org/10.5281/zenodo.10883270" ID-Zenodo-Dep="10883270" httpUri="https://zenodo.org/record/10883270/files/figure.png" pageId="6" pageNumber="1194" startId="6.[159,227,1158,1179]" targetBox="[365,1261,183,1126]" targetPageId="6" targetType="figure">
<paragraph id="8BFE36802273B05924EDEDCAFE19FB7A" blockId="6.[159,1467,1158,1206]" pageId="6" pageNumber="1194">
<emphasis id="B935EA922273B05924EDEDCAFF25FB57" bold="true" box="[159,252,1158,1179]" pageId="6" pageNumber="1194">Figure 2.</emphasis>
<taxonomicName id="4C414D032273B0592560EDCAFD83FB57" authority="(Jaekel, 1894)" baseAuthorityName="Jaekel" baseAuthorityYear="1894" box="[274,602,1158,1179]" class="Elasmobranchii" family="Narcinidae" genus="Titanonarke" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="6" pageNumber="1194" phylum="Chordata" rank="species" species="molini">
<emphasis id="B935EA922273B0592560EDCAFE13FB57" box="[274,458,1158,1179]" italics="true" pageId="6" pageNumber="1194">Titanonarke molini</emphasis>
(
<bibRefCitation id="EFD04B712273B05925AAEDCAFD8BFB57" author="Jaekel, O." box="[472,594,1158,1179]" pageId="6" pageNumber="1194" refId="ref23357" refString="Jaekel, O. 1894. Die eoca ¨ nen Selachier vom Monte Bolca: ein Beitrag zur Morphogenie der Wirbelthiere. Springer, Berlin, 177 pp." type="book" year="1894">Jaekel, 1894</bibRefCitation>
)
</taxonomicName>
from the Eocene Monte Postale site.
<emphasis id="B935EA922273B05927B7EDCAFC20FB57" bold="true" box="[965,1017,1158,1179]" pageId="6" pageNumber="1194">A, B,</emphasis>
MGP-PD 26275/6, holotype in part and counterpart. Scale bars = 100 mm.
</paragraph>
</caption>
<paragraph id="8BFE36802273B0572707EC55FC75FB1D" blockId="6.[845,1467,1305,1934]" lastBlockId="8.[845,1467,891,1296]" lastPageId="8" lastPageNumber="1196" pageId="6" pageNumber="1194">
<emphasis id="B935EA922273B0592707EC55FB9BFAFC" bold="true" box="[885,1090,1305,1328]" pageId="6" pageNumber="1194">Chondrocranium.</emphasis>
The rostral cartilage is expanded, dorsoventrally flattened and trough shaped (
<figureCitation id="137A2A052273B0592130EC75FA53FA9C" box="[1346,1418,1337,1360]" captionStart="Figure 5" captionStartId="8.[159,227,682,703]" captionTargetBox="[365,1255,183,650]" captionTargetId="figure-236@8.[365,1261,183,650]" captionTargetPageId="8" captionText="Figure 5. †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site. A, MCSNV IG.VR.67290, close-up of the head and hyoid apparatus; B, reconstruction. Abbreviations: ao, antorbital cartilage; bb, basibranchials; bbc, basibranchial copula; cb, ceratobranchials; cc, chondrocranium; eb, epibranchials; hb, hypobranchials; hym, hyomandibula; la, labial cartilages; me, Meckels cartilage; nc, nasal capsule; pq, palatoquadrate; ps, pseudohyoid; ra, rostral appendix; rf, rostral fontanelle; ro, rostral cartilage; sca, scapulocoracoid; ss, suprascapula; syn, synarcual. Scale bars = 50 mm." figureDoi="http://doi.org/10.5281/zenodo.10883279" httpUri="https://zenodo.org/record/10883279/files/figure.png" pageId="6" pageNumber="1194">Figs 5</figureCitation>
,
<figureCitation id="137A2A052273B05921ECEC75FA72FA9C" box="[1438,1451,1337,1360]" captionStart="Figure 6" captionStartId="8.[159,227,1806,1827]" captionTargetBox="[365,1252,1347,1774]" captionTargetId="figure-317@8.[365,1261,1347,1774]" captionTargetPageId="8" captionText="Figure 6. †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site. A, the juvenile individual MCSNV IG.VR.91359; B, reconstruction. Abbreviations: ao, antorbital cartilage; cb, ceratobranchials; hym, hyomandibula; la, labial cartilages; me, Meckels cartilage; mes, mesopterygium; met, metapterygium; nc, nasal capsule; pel, prepelvic process; pub, puboischiadic bar; pq, palatoquadrate; pro, propterygium; ps, pseudohyoid; rad, pectoral radials; ro, rostral cartilage; sca, scapulocoracoid; ss, suprascapula; syn, synarcual. Scale bars = 50 mm." figureDoi="http://doi.org/10.5281/zenodo.10883285" httpUri="https://zenodo.org/record/10883285/files/figure.png" pageId="6" pageNumber="1194">6</figureCitation>
), resembling the typical condition of narcinids (Miyake
<emphasis id="B935EA922273B059273FEC35FC59FA5C" box="[845,896,1401,1424]" italics="true" pageId="6" pageNumber="1194">et al</emphasis>
. 1992). The rostrum is long, about one-half of the cranial length, wide anteriorly and tapering proximally towards the nasal capsules. The anterior margin of the rostral cartilage is concave, indicating the presence of a rostral fontanelle (= part of the precerebral fontanelle of Miyake
<emphasis id="B935EA922273B05927DBEF54FC02F9E3" box="[937,987,1560,1583]" italics="true" pageId="6" pageNumber="1194">et al</emphasis>
. 1992), that is present also in
<taxonomicName id="4C414D032273B0592146EF54FA6AF9E3" authorityName="Alcock" authorityYear="1898" box="[1332,1459,1560,1583]" class="Elasmobranchii" family="Narcinidae" genus="Benthobatis" kingdom="Animalia" order="Torpediniformes" pageId="6" pageNumber="1194" phylum="Chordata" rank="genus">
<emphasis id="B935EA922273B0592146EF54FA6AF9E3" box="[1332,1459,1560,1583]" italics="true" pageId="6" pageNumber="1194">Benthobatis</emphasis>
</taxonomicName>
,
<taxonomicName id="4C414D032273B059273FEF74FC67F983" authorityName="Bigelow &amp; Schroeder" authorityYear="1948" box="[845,958,1592,1615]" class="Elasmobranchii" family="Narcinidae" genus="Diplobatis" kingdom="Animalia" order="Torpediniformes" pageId="6" pageNumber="1194" phylum="Chordata" rank="genus">
<emphasis id="B935EA922273B059273FEF74FC67F983" box="[845,958,1592,1615]" italics="true" pageId="6" pageNumber="1194">Diplobatis</emphasis>
</taxonomicName>
,
<taxonomicName id="4C414D032273B05927A2EF74FB99F983" authorityName="Heckel" authorityYear="1846" box="[976,1088,1592,1615]" class="Elasmobranchii" family="Narcinidae" genus="Discopyge" kingdom="Animalia" order="Torpediniformes" pageId="6" pageNumber="1194" phylum="Chordata" rank="genus">
<emphasis id="B935EA922273B05927A2EF74FB99F983" box="[976,1088,1592,1615]" italics="true" pageId="6" pageNumber="1194">Discopyge</emphasis>
</taxonomicName>
and
<taxonomicName id="4C414D032273B059200CEF74FA51F983" authority="(Claeson 2014)" baseAuthorityName="Claeson" baseAuthorityYear="2014" box="[1150,1416,1592,1615]" class="Elasmobranchii" family="Narcinidae" genus="Narcine" kingdom="Animalia" order="Torpediniformes" pageId="6" pageNumber="1194" phylum="Chordata" rank="genus">
<emphasis id="B935EA922273B059200CEF74FB0DF983" box="[1150,1236,1592,1615]" italics="true" pageId="6" pageNumber="1194">Narcine</emphasis>
(
<bibRefCitation id="EFD04B712273B0592095EF74FAA6F983" author="Claeson, K. M." box="[1255,1407,1592,1615]" pageId="6" pageNumber="1194" pagination="597 - 612" refId="ref20593" refString="Claeson, K. M. 2014. The impacts of comparative anatomy of electric rays (Batoidea: Torpediniformes) on their systematic hypotheses. Journal of Morphology, 275, 597 - 612." type="journal article" year="2014">Claeson 2014</bibRefCitation>
)
</taxonomicName>
and which also is supported by the phylogenetic analysis of this study (see below). Small lateral projections (= rostral appendices of
<bibRefCitation id="EFD04B712273B0592799EFD4FB4CF963" author="Holmgren, N." box="[1003,1173,1688,1711]" pageId="6" pageNumber="1194" pagination="1 - 100" refId="ref23007" refString="Holmgren, N. 1941. Studies of the head in fishes. Part 2. Comparative anatomy of the adult selachian skull with remarks on the dorsal fins in sharks. Acta Zoologica, 22, 1 - 100." type="journal article" year="1941">Holmgren 1941</bibRefCitation>
; lateral rostral cartilage of Carvalho 1999) off of the rostral fontanelle were indicated in the
<typeStatus id="54FA88222273B05927E3EF94FC37F923" box="[913,1006,1752,1775]" pageId="6" pageNumber="1194" type="holotype">holotype</typeStatus>
by
<bibRefCitation id="EFD04B712273B0592065EF94FB13F923" author="Carvalho, M. R. de" box="[1047,1226,1752,1775]" pageId="6" pageNumber="1194" refId="ref19921" refString="Carvalho, M. R. de. 2010. Morphology and phylogenetic relationships of the giant electric ray from the Eocene of Monte Bolca, Italy (Chondrichthyes: Torpediniformes). Pp. 183 - 198 in D. K. Elliot, J. G. Maisey, X. Yu &amp; D. Miao (eds) Morphology, phylogeny and paleobiogeography of fossil fishes. Verlag Dr. Friedrich Pfeil, Munich." type="book" year="2010">Carvalho (2010)</bibRefCitation>
. Due to the grout and/ or pigment used on the
<typeStatus id="54FA88222273B0592025EFBBFB6DF8C2" box="[1111,1204,1783,1806]" pageId="6" pageNumber="1194" type="holotype">holotype</typeStatus>
<collectionCode id="ED50AE452273B05920CDEFBBFAFAF8C2" box="[1215,1315,1783,1806]" pageId="6" pageNumber="1194">MGP-PD</collectionCode>
26275/6,
<collectionCode id="ED50AE452273B05921E6EFBBFA63F8C2" box="[1428,1466,1783,1806]" country="Colombia" name="Departamento de Biologia de la Universidad del Valle" pageId="6" pageNumber="1194">UV</collectionCode>
light was not useful to distinguish further details of preserved tissues (see
<figureCitation id="137A2A052273B0592069EE7BFB56F882" box="[1051,1167,1847,1870]" captionStart="Figure 7" captionStartId="9.[159,227,863,884]" captionTargetBox="[317,1309,183,831]" captionTargetId="figure-95@9.[317,1309,183,831]" captionTargetPageId="9" captionText="Figure 7. †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site. A, B, UV images of the holotype MGP-PD 26275/6 showing that it was covered with a pigment reflecting an orange light (see also Supplementary material). The arrows in A indicate a leaf also covered with the pigment, a drop, and some rays that were not distally covered (they are blue/grey, as expected); the arrowhead in A indicates part of the third branch of the antorbital cartilage not covered by pigment; the arrowhead in B indicates the rostral appendix; C, Narcine brasiliensis (TNHC 18512); the arrows indicate the rostral appendices. Scale bars: A, B = 50 mm; C = 10 mm." figureDoi="http://doi.org/10.5281/zenodo.10883287" httpUri="https://zenodo.org/record/10883287/files/figure.png" pageId="6" pageNumber="1194">Fig. 7A, B</figureCitation>
). Ultraviolet light did highlight a possible lateral projection (
<figureCitation id="137A2A052273B05920CFEE1BFAC9F8A2" box="[1213,1296,1879,1902]" captionStart="Figure 7" captionStartId="9.[159,227,863,884]" captionTargetBox="[317,1309,183,831]" captionTargetId="figure-95@9.[317,1309,183,831]" captionTargetPageId="9" captionText="Figure 7. †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site. A, B, UV images of the holotype MGP-PD 26275/6 showing that it was covered with a pigment reflecting an orange light (see also Supplementary material). The arrows in A indicate a leaf also covered with the pigment, a drop, and some rays that were not distally covered (they are blue/grey, as expected); the arrowhead in A indicates part of the third branch of the antorbital cartilage not covered by pigment; the arrowhead in B indicates the rostral appendix; C, Narcine brasiliensis (TNHC 18512); the arrows indicate the rostral appendices. Scale bars: A, B = 50 mm; C = 10 mm." figureDoi="http://doi.org/10.5281/zenodo.10883287" httpUri="https://zenodo.org/record/10883287/files/figure.png" pageId="6" pageNumber="1194">Fig. 7B</figureCitation>
), but we cannot discern whether it was branching and connected to the antorbital cartilage as in
<taxonomicName id="4C414D03227DB05725B9EA37FF07FC7E" authority="(Olfers, 1831)" baseAuthorityName="Olfers" baseAuthorityYear="1831" class="Elasmobranchii" family="Narcinidae" genus="Narcine" kingdom="Animalia" order="Torpediniformes" pageId="8" pageNumber="1196" phylum="Chordata" rank="species" species="brasiliensis">
<emphasis id="B935EA92227DB05725B9EA37FD75FC5E" box="[459,684,891,914]" italics="true" pageId="8" pageNumber="1196">Narcine brasiliensis</emphasis>
(Olfers, 1831)
</taxonomicName>
(
<figureCitation id="137A2A05227DB057249FEAD7FE99FC7E" box="[237,320,923,946]" captionStart="Figure 7" captionStartId="9.[159,227,863,884]" captionTargetBox="[317,1309,183,831]" captionTargetId="figure-95@9.[317,1309,183,831]" captionTargetPageId="9" captionText="Figure 7. †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site. A, B, UV images of the holotype MGP-PD 26275/6 showing that it was covered with a pigment reflecting an orange light (see also Supplementary material). The arrows in A indicate a leaf also covered with the pigment, a drop, and some rays that were not distally covered (they are blue/grey, as expected); the arrowhead in A indicates part of the third branch of the antorbital cartilage not covered by pigment; the arrowhead in B indicates the rostral appendix; C, Narcine brasiliensis (TNHC 18512); the arrows indicate the rostral appendices. Scale bars: A, B = 50 mm; C = 10 mm." figureDoi="http://doi.org/10.5281/zenodo.10883287" httpUri="https://zenodo.org/record/10883287/files/figure.png" pageId="8" pageNumber="1196">Fig. 7C</figureCitation>
). Evidence of a lateral projection off of the rostral fontanelle is, however, clearly present in
<collectionCode id="ED50AE45227DB05726DDEAF7FCD4FC1E" box="[687,781,955,978]" pageId="8" pageNumber="1196">MCSNV</collectionCode>
<collectionCode id="ED50AE45227DB05724EDEA96FF65FC3D" box="[159,188,986,1009]" country="China" name="Institute of Geology" pageId="8" pageNumber="1196">IG</collectionCode>
.VR.67290 (
<figureCitation id="137A2A05227DB0572536EA96FE5FFC3D" box="[324,390,986,1009]" captionStart="Figure 5" captionStartId="8.[159,227,682,703]" captionTargetBox="[365,1255,183,650]" captionTargetId="figure-236@8.[365,1261,183,650]" captionTargetPageId="8" captionText="Figure 5. †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site. A, MCSNV IG.VR.67290, close-up of the head and hyoid apparatus; B, reconstruction. Abbreviations: ao, antorbital cartilage; bb, basibranchials; bbc, basibranchial copula; cb, ceratobranchials; cc, chondrocranium; eb, epibranchials; hb, hypobranchials; hym, hyomandibula; la, labial cartilages; me, Meckels cartilage; nc, nasal capsule; pq, palatoquadrate; ps, pseudohyoid; ra, rostral appendix; rf, rostral fontanelle; ro, rostral cartilage; sca, scapulocoracoid; ss, suprascapula; syn, synarcual. Scale bars = 50 mm." figureDoi="http://doi.org/10.5281/zenodo.10883279" httpUri="https://zenodo.org/record/10883279/files/figure.png" pageId="8" pageNumber="1196">Fig. 5</figureCitation>
), but not in juvenile specimens of †
<taxonomicName id="4C414D03227DB05724D9EAB6FEF4FBDD" authorityName="Carvalho" authorityYear="2010" box="[171,301,1018,1041]" class="Elasmobranchii" family="Narcinidae" genus="Titanonarke" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="8" pageNumber="1196" phylum="Chordata" rank="genus">
<emphasis id="B935EA92227DB05724D9EAB6FEF4FBDD" box="[171,301,1018,1041]" italics="true" pageId="8" pageNumber="1196">Titanonarke</emphasis>
</taxonomicName>
that we examined, which thus resemble juveniles of
<taxonomicName id="4C414D03227DB0572540ED56FDC8FBFD" baseAuthorityName="Olfers" baseAuthorityYear="1831" box="[306,529,1050,1073]" class="Elasmobranchii" family="Narcinidae" genus="Narcine" kingdom="Animalia" order="Torpediniformes" pageId="8" pageNumber="1196" phylum="Chordata" rank="species" species="brasiliensis">
<emphasis id="B935EA92227DB0572540ED56FDC8FBFD" box="[306,529,1050,1073]" italics="true" pageId="8" pageNumber="1196">Narcine brasiliensis</emphasis>
</taxonomicName>
(Miyake
<emphasis id="B935EA92227DB05726F5ED56FD64FBFD" box="[647,701,1050,1073]" italics="true" pageId="8" pageNumber="1196">et al</emphasis>
. 1992, fig. 14A). The connection of the rostral cartilage to the antorbital cartilage through the lateral projection was also detected in
<taxonomicName id="4C414D03227DB057255AED36FE40FB5D" authorityName="Bigelow &amp; Schroeder" authorityYear="1948" box="[296,409,1146,1169]" class="Elasmobranchii" family="Narcinidae" genus="Diplobatis" kingdom="Animalia" order="Torpediniformes" pageId="8" pageNumber="1196" phylum="Chordata" rank="genus">
<emphasis id="B935EA92227DB057255AED36FE40FB5D" box="[296,409,1146,1169]" italics="true" pageId="8" pageNumber="1196">Diplobatis</emphasis>
</taxonomicName>
(
<bibRefCitation id="EFD04B71227DB05725DCED36FCDFFB5D" author="Fechhelm, J. D. &amp; McEachran, J. D." box="[430,774,1146,1169]" pageId="8" pageNumber="1196" pagination="171 - 209" refId="ref21620" refString="Fechhelm, J. D. &amp; McEachran, J. D. 1984. A revision of the electric ray genus Diplobatis with notes on the interrelationships of Narcinidae (Chondrichthyes, Torpediniformes). Bulletin of the Florida State Museum Biological Sciences, 29, 171 - 209." type="journal article" year="1984">Fechhelm &amp; McEachran 1984</bibRefCitation>
, fig. 5),
<taxonomicName id="4C414D03227DB057249EEDD6FEB5FB7D" authorityName="Alcock" authorityYear="1898" box="[236,364,1178,1201]" class="Elasmobranchii" family="Narcinidae" genus="Benthobatis" kingdom="Animalia" order="Torpediniformes" pageId="8" pageNumber="1196" phylum="Chordata" rank="genus">
<emphasis id="B935EA92227DB057249EEDD6FEB5FB7D" box="[236,364,1178,1201]" italics="true" pageId="8" pageNumber="1196">Benthobatis</emphasis>
</taxonomicName>
(Rincon
<emphasis id="B935EA92227DB05725BDEDD6FE27FB7D" box="[463,510,1178,1201]" italics="true" pageId="8" pageNumber="1196">et al</emphasis>
. 2001, fig. 7) and all narcinids in general (
<bibRefCitation id="EFD04B71227DB0572517EDF6FD76FB1D" author="Fechhelm, J. D. &amp; McEachran, J. D." box="[357,687,1210,1233]" pageId="8" pageNumber="1196" pagination="171 - 209" refId="ref21620" refString="Fechhelm, J. D. &amp; McEachran, J. D. 1984. A revision of the electric ray genus Diplobatis with notes on the interrelationships of Narcinidae (Chondrichthyes, Torpediniformes). Bulletin of the Florida State Museum Biological Sciences, 29, 171 - 209." type="journal article" year="1984">Fechhelm &amp; McEachran 1984</bibRefCitation>
, fig. 16; Carvalho 1999, 2010). This character, which is unique among torpediniforms, corroborates the monophyly and the systematic position of narcinids in this study. The juvenile specimen
<collectionCode id="ED50AE45227DB0572064EAD7FBADFC7E" box="[1046,1140,923,946]" pageId="8" pageNumber="1196">MCSNV</collectionCode>
<collectionCode id="ED50AE45227DB057200EEAD7FB40FC7E" box="[1148,1177,923,946]" country="China" name="Institute of Geology" pageId="8" pageNumber="1196">IG</collectionCode>
.VR.91359 shows the early stage of the development of the rostral cartilage (
<figureCitation id="137A2A05227DB0572119EAF7FA72FC1E" box="[1387,1451,955,978]" captionStart="Figure 6" captionStartId="8.[159,227,1806,1827]" captionTargetBox="[365,1252,1347,1774]" captionTargetId="figure-317@8.[365,1261,1347,1774]" captionTargetPageId="8" captionText="Figure 6. †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site. A, the juvenile individual MCSNV IG.VR.91359; B, reconstruction. Abbreviations: ao, antorbital cartilage; cb, ceratobranchials; hym, hyomandibula; la, labial cartilages; me, Meckels cartilage; mes, mesopterygium; met, metapterygium; nc, nasal capsule; pel, prepelvic process; pub, puboischiadic bar; pq, palatoquadrate; pro, propterygium; ps, pseudohyoid; rad, pectoral radials; ro, rostral cartilage; sca, scapulocoracoid; ss, suprascapula; syn, synarcual. Scale bars = 50 mm." figureDoi="http://doi.org/10.5281/zenodo.10883285" httpUri="https://zenodo.org/record/10883285/files/figure.png" pageId="8" pageNumber="1196">Fig. 6</figureCitation>
), with at least one cartilaginous strip (=
<emphasis id="B935EA92227DB057209EEA96FAFEFC3D" box="[1260,1319,986,1009]" italics="true" pageId="8" pageNumber="1196">sensu</emphasis>
Miyake
<emphasis id="B935EA92227DB05721F7EA96FA6DFC3D" box="[1413,1460,986,1009]" italics="true" pageId="8" pageNumber="1196">et al</emphasis>
. 1992) and two lateral rostral bars. The rostral cartilage of †
<taxonomicName id="4C414D03227DB057272AED56FC03FBFD" authorityName="Carvalho" authorityYear="2010" box="[856,986,1050,1073]" class="Elasmobranchii" family="Narcinidae" genus="Titanonarke" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="8" pageNumber="1196" phylum="Chordata" rank="genus">
<emphasis id="B935EA92227DB057272AED56FC03FBFD" box="[856,986,1050,1073]" italics="true" pageId="8" pageNumber="1196">Titanonarke</emphasis>
</taxonomicName>
lacks the basonasal fenestrae that characterize
<taxonomicName id="4C414D03227DB0572707ED76FA90FB9D" authority="(Fechhelm &amp; McEachran 1984)" baseAuthorityName="Fechhelm &amp; McEachran" baseAuthorityYear="1984" box="[885,1353,1082,1105]" class="Elasmobranchii" family="Narcinidae" genus="Diplobatis" kingdom="Animalia" order="Torpediniformes" pageId="8" pageNumber="1196" phylum="Chordata" rank="genus">
<emphasis id="B935EA92227DB0572707ED76FC3FFB9D" box="[885,998,1082,1105]" italics="true" pageId="8" pageNumber="1196">Diplobatis</emphasis>
(
<bibRefCitation id="EFD04B71227DB0572787ED76FAE7FB9D" author="Fechhelm, J. D. &amp; McEachran, J. D." box="[1013,1342,1082,1105]" pageId="8" pageNumber="1196" pagination="171 - 209" refId="ref21620" refString="Fechhelm, J. D. &amp; McEachran, J. D. 1984. A revision of the electric ray genus Diplobatis with notes on the interrelationships of Narcinidae (Chondrichthyes, Torpediniformes). Bulletin of the Florida State Museum Biological Sciences, 29, 171 - 209." type="journal article" year="1984">Fechhelm &amp; McEachran 1984</bibRefCitation>
)
</taxonomicName>
. Although probably present on the dorsal surface of the chondrocranium, it is not possible to distinguish the presence of the anterior and frontoparietal fontanelle in the available material.
</paragraph>
<caption id="DF3E66082272B05824EDEAA1FEACFBD0" ID-DOI="http://doi.org/10.5281/zenodo.10883273" ID-Zenodo-Dep="10883273" httpUri="https://zenodo.org/record/10883273/files/figure.png" pageId="7" pageNumber="1195" startId="7.[159,227,1005,1026]" targetBox="[365,1261,183,973]" targetPageId="7" targetType="figure">
<paragraph id="8BFE36802272B05824EDEAA1FEACFBD0" blockId="7.[159,1467,1004,1053]" pageId="7" pageNumber="1195">
<emphasis id="B935EA922272B05824EDEAA1FF25FBCE" bold="true" box="[159,252,1005,1026]" pageId="7" pageNumber="1195">Figure 3.</emphasis>
<taxonomicName id="4C414D032272B0582560EAA1FD83FBCE" authority="(Jaekel, 1894)" baseAuthorityName="Jaekel" baseAuthorityYear="1894" box="[274,602,1005,1026]" class="Elasmobranchii" family="Narcinidae" genus="Titanonarke" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="7" pageNumber="1195" phylum="Chordata" rank="species" species="molini">
<emphasis id="B935EA922272B0582560EAA1FE12FBCE" box="[274,459,1005,1026]" italics="true" pageId="7" pageNumber="1195">Titanonarke molini</emphasis>
(
<bibRefCitation id="EFD04B712272B05825AAEAA1FD8BFBCE" author="Jaekel, O." box="[472,594,1005,1026]" pageId="7" pageNumber="1195" refId="ref23357" refString="Jaekel, O. 1894. Die eoca ¨ nen Selachier vom Monte Bolca: ein Beitrag zur Morphogenie der Wirbelthiere. Springer, Berlin, 177 pp." type="book" year="1894">Jaekel, 1894</bibRefCitation>
)
</taxonomicName>
from the Eocene Monte Postale site.
<emphasis id="B935EA922272B05827B4EAA1FC07FBCE" bold="true" box="[966,990,1005,1026]" pageId="7" pageNumber="1195">A,</emphasis>
MCSNV IG.VR.67290;
<emphasis id="B935EA922272B05820BDEAA1FB3CFBCE" bold="true" box="[1231,1253,1005,1026]" pageId="7" pageNumber="1195">B,</emphasis>
MCSNV IG.91128/9. Scale bars = 100 mm.
</paragraph>
</caption>
<caption id="DF3E66082272B05824EDEE6CFDEFF89C" ID-DOI="http://doi.org/10.5281/zenodo.10883277" ID-Zenodo-Dep="10883277" httpUri="https://zenodo.org/record/10883277/files/figure.png" pageId="7" pageNumber="1195" startId="7.[159,227,1824,1845]" targetBox="[365,1261,1139,1791]" targetPageId="7" targetType="figure">
<paragraph id="8BFE36802272B05824EDEE6CFDEFF89C" blockId="7.[159,1466,1823,1872]" pageId="7" pageNumber="1195">
<emphasis id="B935EA922272B05824EDEE6CFF22F8F9" bold="true" box="[159,251,1824,1845]" pageId="7" pageNumber="1195">Figure 4.</emphasis>
Juvenile individuals of †
<taxonomicName id="4C414D032272B0582583EE6CFCEEF8F9" authority="(Jaekel, 1894)" baseAuthorityName="Jaekel" baseAuthorityYear="1894" box="[497,823,1824,1845]" class="Elasmobranchii" family="Narcinidae" genus="Titanonarke" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="7" pageNumber="1195" phylum="Chordata" rank="species" species="molini">
<emphasis id="B935EA922272B0582583EE6CFD70F8F9" box="[497,681,1824,1845]" italics="true" pageId="7" pageNumber="1195">Titanonarke molini</emphasis>
(
<bibRefCitation id="EFD04B712272B05826C4EE6CFCF6F8F9" author="Jaekel, O." box="[694,815,1824,1845]" pageId="7" pageNumber="1195" refId="ref23357" refString="Jaekel, O. 1894. Die eoca ¨ nen Selachier vom Monte Bolca: ein Beitrag zur Morphogenie der Wirbelthiere. Springer, Berlin, 177 pp." type="book" year="1894">Jaekel, 1894</bibRefCitation>
)
</taxonomicName>
from the Eocene Monte Postale site.
<emphasis id="B935EA922272B05820ECEE6CFB6CF8F9" bold="true" box="[1182,1205,1824,1845]" pageId="7" pageNumber="1195">A,</emphasis>
MCSNV IG.VR.91359;
<emphasis id="B935EA922272B05821D7EE6CFA63F8F9" bold="true" box="[1445,1466,1824,1845]" pageId="7" pageNumber="1195">B,</emphasis>
MCSNV IG.135581. Scale bars = 10 mm.
</paragraph>
</caption>
<caption id="DF3E6608227DB05724EDEBE6FD7DFCE5" ID-DOI="http://doi.org/10.5281/zenodo.10883279" ID-Zenodo-Dep="10883279" httpUri="https://zenodo.org/record/10883279/files/figure.png" pageId="8" pageNumber="1196" startId="8.[159,227,682,703]" targetBox="[365,1255,183,650]" targetPageId="8" targetType="figure">
<paragraph id="8BFE3680227DB05724EDEBE6FD7DFCE5" blockId="8.[159,1467,681,810]" pageId="8" pageNumber="1196">
<emphasis id="B935EA92227DB05724EDEBE6FF25FD73" bold="true" box="[159,252,682,703]" pageId="8" pageNumber="1196">Figure 5.</emphasis>
<taxonomicName id="4C414D03227DB0572560EBE6FD83FD73" authority="(Jaekel, 1894)" baseAuthorityName="Jaekel" baseAuthorityYear="1894" box="[274,602,682,703]" class="Elasmobranchii" family="Narcinidae" genus="Titanonarke" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="8" pageNumber="1196" phylum="Chordata" rank="species" species="molini">
<emphasis id="B935EA92227DB0572560EBE6FE13FD73" box="[274,458,682,703]" italics="true" pageId="8" pageNumber="1196">Titanonarke molini</emphasis>
(
<bibRefCitation id="EFD04B71227DB05725AAEBE6FD8BFD73" author="Jaekel, O." box="[472,594,682,703]" pageId="8" pageNumber="1196" refId="ref23357" refString="Jaekel, O. 1894. Die eoca ¨ nen Selachier vom Monte Bolca: ein Beitrag zur Morphogenie der Wirbelthiere. Springer, Berlin, 177 pp." type="book" year="1894">Jaekel, 1894</bibRefCitation>
)
</taxonomicName>
from the Eocene Monte Postale site.
<emphasis id="B935EA92227DB05727B4EBE6FC04FD73" bold="true" box="[966,989,682,703]" pageId="8" pageNumber="1196">A,</emphasis>
MCSNV IG.VR.67290, close-up of the head and hyoid apparatus;
<emphasis id="B935EA92227DB0572537EB88FE82FD15" bold="true" box="[325,347,708,729]" pageId="8" pageNumber="1196">B,</emphasis>
reconstruction. Abbreviations: ao, antorbital cartilage; bb, basibranchials; bbc, basibranchial copula; cb, ceratobranchials; cc, chondrocranium; eb, epibranchials; hb, hypobranchials; hym, hyomandibula; la, labial cartilages; me, Meckels cartilage; nc, nasal capsule; pq, palatoquadrate; ps, pseudohyoid; ra, rostral appendix; rf, rostral fontanelle; ro, rostral cartilage; sca, scapulocoracoid; ss, suprascapula; syn, synarcual. Scale bars = 50 mm.
</paragraph>
</caption>
<paragraph id="8BFE3680227DB0552715ED95FA76FBCE" blockId="8.[845,1467,891,1296]" lastBlockId="10.[845,1467,844,1154]" lastPageId="10" lastPageNumber="1198" pageId="8" pageNumber="1196">
The antorbital cartilages in juveniles and adults are robust, well developed, directed anteriorly and broadly branching (
<figureCitation id="137A2A05227CB056256FED56FEB8FBFD" box="[285,353,1050,1073]" captionStart="Figure 5" captionStartId="8.[159,227,682,703]" captionTargetBox="[365,1255,183,650]" captionTargetId="figure-236@8.[365,1261,183,650]" captionTargetPageId="8" captionText="Figure 5. †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site. A, MCSNV IG.VR.67290, close-up of the head and hyoid apparatus; B, reconstruction. Abbreviations: ao, antorbital cartilage; bb, basibranchials; bbc, basibranchial copula; cb, ceratobranchials; cc, chondrocranium; eb, epibranchials; hb, hypobranchials; hym, hyomandibula; la, labial cartilages; me, Meckels cartilage; nc, nasal capsule; pq, palatoquadrate; ps, pseudohyoid; ra, rostral appendix; rf, rostral fontanelle; ro, rostral cartilage; sca, scapulocoracoid; ss, suprascapula; syn, synarcual. Scale bars = 50 mm." figureDoi="http://doi.org/10.5281/zenodo.10883279" httpUri="https://zenodo.org/record/10883279/files/figure.png" pageId="9" pageNumber="1197">Figs 5</figureCitation>
,
<figureCitation id="137A2A05227CB0562500ED56FEA6FBFD" box="[370,383,1050,1073]" captionStart="Figure 6" captionStartId="8.[159,227,1806,1827]" captionTargetBox="[365,1252,1347,1774]" captionTargetId="figure-317@8.[365,1261,1347,1774]" captionTargetPageId="8" captionText="Figure 6. †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site. A, the juvenile individual MCSNV IG.VR.91359; B, reconstruction. Abbreviations: ao, antorbital cartilage; cb, ceratobranchials; hym, hyomandibula; la, labial cartilages; me, Meckels cartilage; mes, mesopterygium; met, metapterygium; nc, nasal capsule; pel, prepelvic process; pub, puboischiadic bar; pq, palatoquadrate; pro, propterygium; ps, pseudohyoid; rad, pectoral radials; ro, rostral cartilage; sca, scapulocoracoid; ss, suprascapula; syn, synarcual. Scale bars = 50 mm." figureDoi="http://doi.org/10.5281/zenodo.10883285" httpUri="https://zenodo.org/record/10883285/files/figure.png" pageId="9" pageNumber="1197">6</figureCitation>
). There are no distinct foramina on the anterior cartilages. Antorbitals are subdivided at about one-third of their length into two main branches; their stout bases articulate with the lateral aspect of the nasal capsules through an expanded posterior condyle. The distal end of the largest branch is broadly separated from the anterior extension of the propterygium. A third, smaller, posteriorly directed branch is located at midlength of the antorbital cartilages (
<figureCitation id="137A2A05227CB05620B3ED36FADAFB5D" box="[1217,1283,1146,1169]" captionStart="Figure 5" captionStartId="8.[159,227,682,703]" captionTargetBox="[365,1255,183,650]" captionTargetId="figure-236@8.[365,1261,183,650]" captionTargetPageId="8" captionText="Figure 5. †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site. A, MCSNV IG.VR.67290, close-up of the head and hyoid apparatus; B, reconstruction. Abbreviations: ao, antorbital cartilage; bb, basibranchials; bbc, basibranchial copula; cb, ceratobranchials; cc, chondrocranium; eb, epibranchials; hb, hypobranchials; hym, hyomandibula; la, labial cartilages; me, Meckels cartilage; nc, nasal capsule; pq, palatoquadrate; ps, pseudohyoid; ra, rostral appendix; rf, rostral fontanelle; ro, rostral cartilage; sca, scapulocoracoid; ss, suprascapula; syn, synarcual. Scale bars = 50 mm." figureDoi="http://doi.org/10.5281/zenodo.10883279" httpUri="https://zenodo.org/record/10883279/files/figure.png" pageId="9" pageNumber="1197">Figs 5</figureCitation>
,
<figureCitation id="137A2A05227CB0562163ED36FAC7FB5D" box="[1297,1310,1146,1169]" captionStart="Figure 6" captionStartId="8.[159,227,1806,1827]" captionTargetBox="[365,1252,1347,1774]" captionTargetId="figure-317@8.[365,1261,1347,1774]" captionTargetPageId="8" captionText="Figure 6. †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site. A, the juvenile individual MCSNV IG.VR.91359; B, reconstruction. Abbreviations: ao, antorbital cartilage; cb, ceratobranchials; hym, hyomandibula; la, labial cartilages; me, Meckels cartilage; mes, mesopterygium; met, metapterygium; nc, nasal capsule; pel, prepelvic process; pub, puboischiadic bar; pq, palatoquadrate; pro, propterygium; ps, pseudohyoid; rad, pectoral radials; ro, rostral cartilage; sca, scapulocoracoid; ss, suprascapula; syn, synarcual. Scale bars = 50 mm." figureDoi="http://doi.org/10.5281/zenodo.10883285" httpUri="https://zenodo.org/record/10883285/files/figure.png" pageId="9" pageNumber="1197">6</figureCitation>
). The absence of the posterior branch was considered diagnostic for †
<taxonomicName id="4C414D03227FB05524D9EA00FEF4FCAF" authorityName="Carvalho" authorityYear="2010" box="[171,301,844,867]" class="Elasmobranchii" family="Narcinidae" genus="Titanonarke" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="10" pageNumber="1198" phylum="Chordata" rank="genus">
<emphasis id="B935EA92227FB05524D9EA00FEF4FCAF" box="[171,301,844,867]" italics="true" pageId="10" pageNumber="1198">Titanonarke</emphasis>
</taxonomicName>
to the exclusion of
<taxonomicName id="4C414D03227FB055267CEA00FDBDFCAF" baseAuthorityName="Claeson" baseAuthorityYear="2014" box="[526,612,844,867]" class="Elasmobranchii" family="Narcinidae" genus="Narcine" kingdom="Animalia" order="Torpediniformes" pageId="10" pageNumber="1198" phylum="Chordata" rank="genus">
<emphasis id="B935EA92227FB055267CEA00FDBDFCAF" box="[526,612,844,867]" italics="true" pageId="10" pageNumber="1198">Narcine</emphasis>
</taxonomicName>
and
<taxonomicName id="4C414D03227FB05526EFEA00FCD4FCAF" authorityName="Heckel" authorityYear="1846" box="[669,781,844,867]" class="Elasmobranchii" family="Narcinidae" genus="Discopyge" kingdom="Animalia" order="Torpediniformes" pageId="10" pageNumber="1198" phylum="Chordata" rank="genus">
<emphasis id="B935EA92227FB05526EFEA00FCD4FCAF" box="[669,781,844,867]" italics="true" pageId="10" pageNumber="1198">Discopyge</emphasis>
</taxonomicName>
by
<bibRefCitation id="EFD04B71227FB05524B0EA20FEACFC4F" author="Carvalho, M. R. de" box="[194,373,876,899]" pageId="10" pageNumber="1198" refId="ref19921" refString="Carvalho, M. R. de. 2010. Morphology and phylogenetic relationships of the giant electric ray from the Eocene of Monte Bolca, Italy (Chondrichthyes: Torpediniformes). Pp. 183 - 198 in D. K. Elliot, J. G. Maisey, X. Yu &amp; D. Miao (eds) Morphology, phylogeny and paleobiogeography of fossil fishes. Verlag Dr. Friedrich Pfeil, Munich." type="book" year="2010">Carvalho (2010)</bibRefCitation>
. However, the presence of this posterior branch in
<collectionCode id="ED50AE45227FB055252FEAC0FE62FC6F" box="[349,443,908,931]" pageId="10" pageNumber="1198">MCSNV</collectionCode>
<collectionCode id="ED50AE45227FB05525BCEAC0FE35FC6F" box="[462,492,908,931]" country="China" name="Institute of Geology" pageId="10" pageNumber="1198">IG</collectionCode>
.VR.67290,
<collectionCode id="ED50AE45227FB055260BEAC0FD0EFC6F" box="[633,727,908,931]" pageId="10" pageNumber="1198">MCSNV</collectionCode>
<collectionCode id="ED50AE45227FB0552698EAC0FCD0FC6F" box="[746,777,908,931]" country="China" name="Institute of Geology" pageId="10" pageNumber="1198">IG</collectionCode>
. VR.91359,
<collectionCode id="ED50AE45227FB0552569EAE0FEA0FC0F" box="[283,377,940,963]" pageId="10" pageNumber="1198">MCSNV</collectionCode>
<collectionCode id="ED50AE45227FB05525F3EAE0FE46FC0F" box="[385,415,940,963]" country="China" name="Institute of Geology" pageId="10" pageNumber="1198">IG</collectionCode>
.135581 and possibly, at least partially, in the
<typeStatus id="54FA8822227FB055255CEA80FE52FC2F" box="[302,395,972,995]" pageId="10" pageNumber="1198" type="holotype">holotype</typeStatus>
(
<figureCitation id="137A2A05227FB05525EFEA80FE2EFC2F" box="[413,503,972,995]" captionStart="Figure 7" captionStartId="9.[159,227,863,884]" captionTargetBox="[317,1309,183,831]" captionTargetId="figure-95@9.[317,1309,183,831]" captionTargetPageId="9" captionText="Figure 7. †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site. A, B, UV images of the holotype MGP-PD 26275/6 showing that it was covered with a pigment reflecting an orange light (see also Supplementary material). The arrows in A indicate a leaf also covered with the pigment, a drop, and some rays that were not distally covered (they are blue/grey, as expected); the arrowhead in A indicates part of the third branch of the antorbital cartilage not covered by pigment; the arrowhead in B indicates the rostral appendix; C, Narcine brasiliensis (TNHC 18512); the arrows indicate the rostral appendices. Scale bars: A, B = 50 mm; C = 10 mm." figureDoi="http://doi.org/10.5281/zenodo.10883287" httpUri="https://zenodo.org/record/10883287/files/figure.png" pageId="10" pageNumber="1198">Figs 7A</figureCitation>
), suggests that it was not discernible in the specimens described by
<bibRefCitation id="EFD04B71227FB05526D9EAA7FF31FBEE" author="Carvalho, M. R. de" pageId="10" pageNumber="1198" refId="ref19921" refString="Carvalho, M. R. de. 2010. Morphology and phylogenetic relationships of the giant electric ray from the Eocene of Monte Bolca, Italy (Chondrichthyes: Torpediniformes). Pp. 183 - 198 in D. K. Elliot, J. G. Maisey, X. Yu &amp; D. Miao (eds) Morphology, phylogeny and paleobiogeography of fossil fishes. Verlag Dr. Friedrich Pfeil, Munich." type="book" year="2010">Carvalho (2010)</bibRefCitation>
. Moreover, since osteological, morphometric and meristic features are not useful to separate juvenile specimens from the
<typeStatus id="54FA8822227FB0552534ED07FE7FFBAE" box="[326,422,1099,1122]" pageId="10" pageNumber="1198" type="holotype">holotype</typeStatus>
, there is no reason to assign this smaller specimens to a new genus, if only based on the presence of this character. In fact, the
<collectionCode id="ED50AE45227FB0552080EA00FAFCFCAF" box="[1266,1317,844,867]" pageId="10" pageNumber="1198">PCA</collectionCode>
performed in this study on the entire morphological data set of standardized and log-transformed measurements and counts (
<figureCitation id="137A2A05227FB0552724EAE0FC4EFC0F" box="[854,919,940,963]" captionStart="Figure 8" captionStartId="9.[159,227,1833,1854]" captionTargetBox="[318,1308,1256,1800]" captionTargetId="figure-211@9.[317,1309,1255,1801]" captionTargetPageId="9" captionText="Figure 8. Visual image of the principal component analysis (PCA) performed on the entire set of log-transformed standardized morphometric and meristic features, showing the separation of the specimens referred to †Titanonarke molini (right side of morphospace) from those referred to †Titanonarke megapterygia sp. nov. (left side). The illustrations lying along the extreme values of PC1 represent the hypothetical reconstruction of the two species based on body proportions." figureDoi="http://doi.org/10.5281/zenodo.10883289" httpUri="https://zenodo.org/record/10883289/files/figure.png" pageId="10" pageNumber="1198">Fig. 8</figureCitation>
) shows no remarkable separation of the juvenile specimens
<collectionCode id="ED50AE45227FB05527ADEA80FBE4FC2F" box="[991,1085,972,995]" pageId="10" pageNumber="1198">MCSNV</collectionCode>
<collectionCode id="ED50AE45227FB0552012EA80FBA4FC2F" box="[1120,1149,972,995]" country="China" name="Institute of Geology" pageId="10" pageNumber="1198">IG</collectionCode>
.VR.91359 and
<collectionCode id="ED50AE45227FB055212EEA80FA63FC2F" box="[1372,1466,972,995]" pageId="10" pageNumber="1198">MCSNV</collectionCode>
<collectionCode id="ED50AE45227FB055273FEAA7FCB2FBCE" box="[845,875,1003,1026]" country="China" name="Institute of Geology" pageId="10" pageNumber="1198">IG</collectionCode>
.135581 from the
<typeStatus id="54FA8822227FB0552055EAA7FB5DFBCE" box="[1063,1156,1003,1026]" pageId="10" pageNumber="1198" type="holotype">holotype</typeStatus>
and other referred material.
</paragraph>
<caption id="DF3E6608227DB05724EDEE42FEB0F842" ID-DOI="http://doi.org/10.5281/zenodo.10883285" ID-Zenodo-Dep="10883285" httpUri="https://zenodo.org/record/10883285/files/figure.png" pageId="8" pageNumber="1196" startId="8.[159,227,1806,1827]" targetBox="[365,1252,1347,1774]" targetPageId="8" targetType="figure">
<paragraph id="8BFE3680227DB05724EDEE42FEB0F842" blockId="8.[159,1467,1806,1934]" pageId="8" pageNumber="1196">
<emphasis id="B935EA92227DB05724EDEE42FF22F8EF" bold="true" box="[159,251,1806,1827]" pageId="8" pageNumber="1196">Figure 6.</emphasis>
<taxonomicName id="4C414D03227DB0572563EE42FD8EF8EF" authority="(Jaekel, 1894)" baseAuthorityName="Jaekel" baseAuthorityYear="1894" box="[273,599,1806,1827]" class="Elasmobranchii" family="Narcinidae" genus="Titanonarke" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="8" pageNumber="1196" phylum="Chordata" rank="species" species="molini">
<emphasis id="B935EA92227DB0572563EE42FE10F8EF" box="[273,457,1806,1827]" italics="true" pageId="8" pageNumber="1196">Titanonarke molini</emphasis>
(
<bibRefCitation id="EFD04B71227DB05725A4EE42FD89F8EF" author="Jaekel, O." box="[470,592,1806,1827]" pageId="8" pageNumber="1196" refId="ref23357" refString="Jaekel, O. 1894. Die eoca ¨ nen Selachier vom Monte Bolca: ein Beitrag zur Morphogenie der Wirbelthiere. Springer, Berlin, 177 pp." type="book" year="1894">Jaekel, 1894</bibRefCitation>
)
</taxonomicName>
from the Eocene Monte Postale site.
<emphasis id="B935EA92227DB05727B2EE42FC0EF8EF" bold="true" box="[960,983,1806,1827]" pageId="8" pageNumber="1196">A,</emphasis>
the juvenile individual MCSNV IG.VR.91359;
<emphasis id="B935EA92227DB05721D7EE42FA63F8EF" bold="true" box="[1445,1466,1806,1827]" pageId="8" pageNumber="1196">B,</emphasis>
reconstruction. Abbreviations: ao, antorbital cartilage; cb, ceratobranchials; hym, hyomandibula; la, labial cartilages; me, Meckels cartilage; mes, mesopterygium; met, metapterygium; nc, nasal capsule; pel, prepelvic process; pub, puboischiadic bar; pq, palatoquadrate; pro, propterygium; ps, pseudohyoid; rad, pectoral radials; ro, rostral cartilage; sca, scapulocoracoid; ss, suprascapula; syn, synarcual. Scale bars = 50 mm.
</paragraph>
</caption>
<caption id="DF3E6608227CB05624EDEA13FAE2FC12" ID-DOI="http://doi.org/10.5281/zenodo.10883287" ID-Zenodo-Dep="10883287" httpUri="https://zenodo.org/record/10883287/files/figure.png" pageId="9" pageNumber="1197" startId="9.[159,227,863,884]" targetBox="[317,1309,183,831]" targetPageId="9" targetType="figure">
<paragraph id="8BFE3680227CB05624EDEA13FAE2FC12" blockId="9.[159,1467,862,990]" pageId="9" pageNumber="1197">
<emphasis id="B935EA92227CB05624EDEA13FF25FCB8" bold="true" box="[159,252,863,884]" pageId="9" pageNumber="1197">Figure 7.</emphasis>
<taxonomicName id="4C414D03227CB0562560EA13FD83FCB8" authority="(Jaekel, 1894)" baseAuthorityName="Jaekel" baseAuthorityYear="1894" box="[274,602,863,884]" class="Elasmobranchii" family="Narcinidae" genus="Titanonarke" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="9" pageNumber="1197" phylum="Chordata" rank="species" species="molini">
<emphasis id="B935EA92227CB0562560EA13FE13FCB8" box="[274,458,863,884]" italics="true" pageId="9" pageNumber="1197">Titanonarke molini</emphasis>
(
<bibRefCitation id="EFD04B71227CB05625AAEA13FD8BFCB8" author="Jaekel, O." box="[472,594,863,884]" pageId="9" pageNumber="1197" refId="ref23357" refString="Jaekel, O. 1894. Die eoca ¨ nen Selachier vom Monte Bolca: ein Beitrag zur Morphogenie der Wirbelthiere. Springer, Berlin, 177 pp." type="book" year="1894">Jaekel, 1894</bibRefCitation>
)
</taxonomicName>
from the Eocene Monte Postale site.
<emphasis id="B935EA92227CB05627B7EA13FC20FCB8" bold="true" box="[965,1017,863,884]" pageId="9" pageNumber="1197">A, B,</emphasis>
UV images of the holotype MGP-PD 26275/6 showing that it was covered with a pigment reflecting an orange light (see also Supplementary material). The arrows in A indicate a leaf also covered with the pigment, a drop, and some rays that were not distally covered (they are blue/grey, as expected); the arrowhead in A indicates part of the third branch of the antorbital cartilage not covered by pigment; the arrowhead in B indicates the rostral appendix;
<emphasis id="B935EA92227CB05624EDEA85FF6FFC12" bold="true" box="[159,182,969,990]" pageId="9" pageNumber="1197">C,</emphasis>
<taxonomicName id="4C414D03227CB05624CEEA85FEA4FC12" baseAuthorityName="Olfers" baseAuthorityYear="1831" box="[188,381,969,990]" class="Elasmobranchii" family="Narcinidae" genus="Narcine" kingdom="Animalia" order="Torpediniformes" pageId="9" pageNumber="1197" phylum="Chordata" rank="species" species="brasiliensis">
<emphasis id="B935EA92227CB05624CEEA85FEA4FC12" box="[188,381,969,990]" italics="true" pageId="9" pageNumber="1197">Narcine brasiliensis</emphasis>
</taxonomicName>
(TNHC 18512); the arrows indicate the rostral appendices. Scale bars: A, B = 50 mm; C = 10 mm.
</paragraph>
</caption>
<caption id="DF3E6608227CB05624EDEE65FCB8F842" ID-DOI="http://doi.org/10.5281/zenodo.10883289" ID-Zenodo-Dep="10883289" httpUri="https://zenodo.org/record/10883289/files/figure.png" pageId="9" pageNumber="1197" startId="9.[159,227,1833,1854]" targetBox="[318,1308,1256,1800]" targetPageId="9" targetType="figure">
<paragraph id="8BFE3680227CB05624EDEE65FCB8F842" blockId="9.[159,1467,1833,1934]" pageId="9" pageNumber="1197">
<emphasis id="B935EA92227CB05624EDEE65FF22F8F2" bold="true" box="[159,251,1833,1854]" pageId="9" pageNumber="1197">Figure 8.</emphasis>
Visual image of the principal component analysis (PCA) performed on the entire set of log-transformed standardized morphometric and meristic features, showing the separation of the specimens referred to †
<taxonomicName id="4C414D03227CB05627C9EE08FBADF895" baseAuthorityName="Jaekel" baseAuthorityYear="1894" box="[955,1140,1860,1881]" class="Elasmobranchii" family="Narcinidae" genus="Titanonarke" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="9" pageNumber="1197" phylum="Chordata" rank="species" species="molini">
<emphasis id="B935EA92227CB05627C9EE08FBADF895" box="[955,1140,1860,1881]" italics="true" pageId="9" pageNumber="1197">Titanonarke molini</emphasis>
</taxonomicName>
(right side of morphospace) from those referred to †
<taxonomicName id="4C414D03227CB0562523EE12FD8BF8BF" authority="Marram̀ &amp; Claeso &amp; Carneval &amp; Kriwe, 2018" authorityName="Marram̀ &amp; Claeso &amp; Carneval &amp; Kriwe" authorityYear="2018" box="[337,594,1886,1907]" class="Elasmobranchii" family="Narcinidae" genus="Titanonarke" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="9" pageNumber="1197" phylum="Chordata" rank="species" species="megapterygia" status="sp. nov.">
<emphasis id="B935EA92227CB0562523EE12FD8BF8BF" box="[337,594,1886,1907]" italics="true" pageId="9" pageNumber="1197">Titanonarke megapterygia</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A20657E9227CB056262BEE12FD7FF8BF" box="[601,678,1886,1907]" pageId="9" pageNumber="1197" rank="species">sp. nov.</taxonomicNameLabel>
(left side). The illustrations lying along the extreme values of PC1 represent the hypothetical reconstruction of the two species based on body proportions.
</paragraph>
</caption>
<caption id="DF3E6608227FB05524EDEBF2FCEEFCEF" ID-DOI="http://doi.org/10.5281/zenodo.10883293" ID-Zenodo-Dep="10883293" httpUri="https://zenodo.org/record/10883293/files/figure.png" pageId="10" pageNumber="1198" startId="10.[159,227,702,723]" targetBox="[365,1261,183,670]" targetPageId="10" targetType="figure">
<paragraph id="8BFE3680227FB05524EDEBF2FCEEFCEF" blockId="10.[159,1467,702,804]" pageId="10" pageNumber="1198">
<emphasis id="B935EA92227FB05524EDEBF2FEC6FD1F" bold="true" box="[159,287,702,723]" pageId="10" pageNumber="1198">Figure 9. A,</emphasis>
synarcual and pectoral girdle of
<taxonomicName id="4C414D03227FB0552611EBF2FCFCFD1F" baseAuthorityName="Olfers" baseAuthorityYear="1831" box="[611,805,702,723]" class="Elasmobranchii" family="Narcinidae" genus="Narcine" kingdom="Animalia" order="Torpediniformes" pageId="10" pageNumber="1198" phylum="Chordata" rank="species" species="brasiliensis">
<emphasis id="B935EA92227FB0552611EBF2FCFCFD1F" box="[611,805,702,723]" italics="true" pageId="10" pageNumber="1198">Narcine brasiliensis</emphasis>
</taxonomicName>
(AMNH 95343) in dorsal view;
<emphasis id="B935EA92227FB055201AEBF2FBA7FD1F" bold="true" box="[1128,1150,702,723]" pageId="10" pageNumber="1198">B,</emphasis>
synarcual and pectoral girdle of †
<taxonomicName id="4C414D03227FB05524D8EB95FEBAFD22" baseAuthorityName="Jaekel" baseAuthorityYear="1894" box="[170,355,729,750]" class="Elasmobranchii" family="Narcinidae" genus="Titanonarke" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="10" pageNumber="1198" phylum="Chordata" rank="species" species="molini">
<emphasis id="B935EA92227FB05524D8EB95FEBAFD22" box="[170,355,729,750]" italics="true" pageId="10" pageNumber="1198">Titanonarke molini</emphasis>
</taxonomicName>
(MCSNV IG.VR.91359) in ventral view. The arrowheads indicate the posteriorly directed lateral stays of the synarcual. The arrows indicate the posteriorly directed scapular process of the scapulocoracoid. Abbreviations: cb, ceratocranchials; sca, scapulocoracoid; ss, suprascapula; syn, synarcual. Scale bar = 5 mm.
</paragraph>
</caption>
<paragraph id="8BFE3680227FB0542715ED47FBC7FC7C" blockId="10.[845,1467,844,1154]" lastBlockId="11.[845,1466,826,944]" lastPageId="11" lastPageNumber="1199" pageId="10" pageNumber="1198">
The nasal capsules are ovoid in shape, located at about midlength of the chondrocranium and projected laterally. The internasal plate between the two nasal capsules appears flat and narrow. As all specimens are preserved in dorsoventral view, it was not possible to detect the lateral aspect of the nasal capsules. The otic capsules and the posterior part of the chondrocranium are partially hidden by the jaws and hyoid arches. However, they appear to be approximately as wide as the widest part of the rostrum and much narrower than the nasal capsules. A large basicranial fenestra is recognizable in the juvenile
<collectionCode id="ED50AE45227EB05426DDEAB5FCD4FBDC" box="[687,781,1017,1040]" pageId="11" pageNumber="1199">MCSNV</collectionCode>
<collectionCode id="ED50AE45227EB054273FEA76FCB2FC9D" box="[845,875,826,849]" country="China" name="Institute of Geology" pageId="11" pageNumber="1199">IG</collectionCode>
.VR.91359 (
<figureCitation id="137A2A05227EB0542780EA76FBEDFC9D" box="[1010,1076,826,849]" captionStart="Figure 6" captionStartId="8.[159,227,1806,1827]" captionTargetBox="[365,1252,1347,1774]" captionTargetId="figure-317@8.[365,1261,1347,1774]" captionTargetPageId="8" captionText="Figure 6. †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site. A, the juvenile individual MCSNV IG.VR.91359; B, reconstruction. Abbreviations: ao, antorbital cartilage; cb, ceratobranchials; hym, hyomandibula; la, labial cartilages; me, Meckels cartilage; mes, mesopterygium; met, metapterygium; nc, nasal capsule; pel, prepelvic process; pub, puboischiadic bar; pq, palatoquadrate; pro, propterygium; ps, pseudohyoid; rad, pectoral radials; ro, rostral cartilage; sca, scapulocoracoid; ss, suprascapula; syn, synarcual. Scale bars = 50 mm." figureDoi="http://doi.org/10.5281/zenodo.10883285" httpUri="https://zenodo.org/record/10883285/files/figure.png" pageId="11" pageNumber="1199">Fig. 6</figureCitation>
), which resembles in position and shape the condition shown in the early stages of chondrification of the otic region in
<taxonomicName id="4C414D03227EB054200FEA35FA8EFC5C" baseAuthorityName="Olfers" baseAuthorityYear="1831" box="[1149,1367,889,912]" class="Elasmobranchii" family="Narcinidae" genus="Narcine" kingdom="Animalia" order="Torpediniformes" pageId="11" pageNumber="1199" phylum="Chordata" rank="species" species="brasiliensis">
<emphasis id="B935EA92227EB054200FEA35FA8EFC5C" box="[1149,1367,889,912]" italics="true" pageId="11" pageNumber="1199">Narcine brasiliensis</emphasis>
</taxonomicName>
(Miyake
<emphasis id="B935EA92227EB054273FEAD5FCA5FC7C" box="[845,892,921,944]" italics="true" pageId="11" pageNumber="1199">et al</emphasis>
. 1992, fig. 14).
</paragraph>
<caption id="DF3E6608227FB05524EDEE65FEF6F842" ID-DOI="http://doi.org/10.5281/zenodo.10883295" ID-Zenodo-Dep="10883295" httpUri="https://zenodo.org/record/10883295/files/figure.png" pageId="10" pageNumber="1198" startId="10.[159,227,1833,1854]" targetBox="[365,1248,1190,1801]" targetPageId="10" targetType="figure">
<paragraph id="8BFE3680227FB05524EDEE65FEF6F842" blockId="10.[159,1467,1832,1934]" pageId="10" pageNumber="1198">
<emphasis id="B935EA92227FB05524EDEE65FED1F8F2" bold="true" box="[159,264,1833,1854]" pageId="10" pageNumber="1198">Figure 10.</emphasis>
Pectoral fin of †
<taxonomicName id="4C414D03227FB05525D9EE65FD2BF8F2" authority="(Jaekel, 1894)" baseAuthorityName="Jaekel" baseAuthorityYear="1894" box="[427,754,1833,1854]" class="Elasmobranchii" family="Narcinidae" genus="Titanonarke" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="10" pageNumber="1198" phylum="Chordata" rank="species" species="molini">
<emphasis id="B935EA92227FB05525D9EE65FDBDF8F2" box="[427,612,1833,1854]" italics="true" pageId="10" pageNumber="1198">Titanonarke molini</emphasis>
(
<bibRefCitation id="EFD04B71227FB0552602EE65FD33F8F2" author="Jaekel, O." box="[624,746,1833,1854]" pageId="10" pageNumber="1198" refId="ref23357" refString="Jaekel, O. 1894. Die eoca ¨ nen Selachier vom Monte Bolca: ein Beitrag zur Morphogenie der Wirbelthiere. Springer, Berlin, 177 pp." type="book" year="1894">Jaekel, 1894</bibRefCitation>
)
</taxonomicName>
from the Eocene Monte Postale site.
<emphasis id="B935EA92227FB0552028EE65FBABF8F2" bold="true" box="[1114,1138,1833,1854]" pageId="10" pageNumber="1198">A,</emphasis>
MCSNV IG.VR.67290;
<emphasis id="B935EA92227FB0552113EE65FAAEF8F2" bold="true" box="[1377,1399,1833,1854]" pageId="10" pageNumber="1198">B,</emphasis>
reconstruction; different colours are used to distinguish the propterygial (green), mesopterygial (yellow) and metapterygial (red) radials (colours in the online version). Abbreviations: mes, mesopterygium; met, metapterygium; pro, propterygium; sca, scapulocoracoid. Scale bars = 50 mm.
</paragraph>
</caption>
<caption id="DF3E6608227EB05424EDEBF1FD0CFCCB" ID-DOI="http://doi.org/10.5281/zenodo.10883297" ID-Zenodo-Dep="10883297" httpUri="https://zenodo.org/record/10883297/files/figure.png" pageId="11" pageNumber="1199" startId="11.[159,227,701,722]" targetBox="[365,1257,183,669]" targetPageId="11" targetType="figure">
<paragraph id="8BFE3680227EB05424EDEBF1FD0CFCCB" blockId="11.[159,1466,701,776]" pageId="11" pageNumber="1199">
<emphasis id="B935EA92227EB05424EDEBF1FED1FD1E" bold="true" box="[159,264,701,722]" pageId="11" pageNumber="1199">Figure 11.</emphasis>
Pelvic fins of †
<taxonomicName id="4C414D03227EB05425D6EBF1FD35FD1E" authority="(Jaekel, 1894)" baseAuthorityName="Jaekel" baseAuthorityYear="1894" box="[420,748,701,722]" class="Elasmobranchii" family="Narcinidae" genus="Titanonarke" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="11" pageNumber="1199" phylum="Chordata" rank="species" species="molini">
<emphasis id="B935EA92227EB05425D6EBF1FD84FD1E" box="[420,605,701,722]" italics="true" pageId="11" pageNumber="1199">Titanonarke molini</emphasis>
(
<bibRefCitation id="EFD04B71227EB0542618EBF1FD3DFD1E" author="Jaekel, O." box="[618,740,701,722]" pageId="11" pageNumber="1199" refId="ref23357" refString="Jaekel, O. 1894. Die eoca ¨ nen Selachier vom Monte Bolca: ein Beitrag zur Morphogenie der Wirbelthiere. Springer, Berlin, 177 pp." type="book" year="1894">Jaekel, 1894</bibRefCitation>
)
</taxonomicName>
from the Eocene Monte Postale site.
<emphasis id="B935EA92227EB054202AEBF1FBB6FD1E" bold="true" box="[1112,1135,701,722]" pageId="11" pageNumber="1199">A,</emphasis>
MCSNV IG.VR.67290;
<emphasis id="B935EA92227EB0542113EBF1FAAFFD1E" bold="true" box="[1377,1398,701,722]" pageId="11" pageNumber="1199">B,</emphasis>
reconstruction. Abbreviations: bas, basipterygia; ilp, iliac process; pel, pelvic processes; pub, puboischiadic bar; rad, pelvic radials. The arrowhead indicates the iliac process. Scale bars = 50 mm.
</paragraph>
</caption>
<paragraph id="8BFE3680227EB0542707EA86FB6AF912" blockId="11.[845,1467,970,1758]" pageId="11" pageNumber="1199">
<emphasis id="B935EA92227EB0542707EA86FC6DFC2D" bold="true" box="[885,948,970,993]" pageId="11" pageNumber="1199">Jaws.</emphasis>
<bibRefCitation id="EFD04B71227EB05427B0EA86FBA3FC2D" author="Carvalho, M. R. de" box="[962,1146,970,993]" pageId="11" pageNumber="1199" refId="ref19921" refString="Carvalho, M. R. de. 2010. Morphology and phylogenetic relationships of the giant electric ray from the Eocene of Monte Bolca, Italy (Chondrichthyes: Torpediniformes). Pp. 183 - 198 in D. K. Elliot, J. G. Maisey, X. Yu &amp; D. Miao (eds) Morphology, phylogeny and paleobiogeography of fossil fishes. Verlag Dr. Friedrich Pfeil, Munich." type="book" year="2010">Carvalho (2010)</bibRefCitation>
considered both palatoquadrate and Meckels cartilage of †
<taxonomicName id="4C414D03227EB05420C3EAA6FACBFBCD" baseAuthorityName="Jaekel" baseAuthorityYear="1894" box="[1201,1298,1002,1025]" class="Elasmobranchii" family="Narcinidae" genus="Titanonarke" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="11" pageNumber="1199" phylum="Chordata" rank="species" species="molini">
<emphasis id="B935EA92227EB05420C3EAA6FACBFBCD" box="[1201,1298,1002,1025]" italics="true" pageId="11" pageNumber="1199">T. molini</emphasis>
</taxonomicName>
broadly arched and not very stout when compared to those of other members of
<taxonomicName id="4C414D03227EB05427D6ED65FBC4FB8C" authorityName="Gill" authorityYear="1862" box="[932,1053,1065,1088]" class="Elasmobranchii" family="Narcinidae" kingdom="Animalia" order="Torpediniformes" pageId="11" pageNumber="1199" phylum="Chordata" rank="family">Narcinidae</taxonomicName>
. Given that long, slender and curved jaws are diagnostic for the torpedinoids
<taxonomicName id="4C414D03227EB054208AED05FA8BFBAC" authorityName="Houttuyn" authorityYear="1764" box="[1272,1362,1097,1120]" class="Elasmobranchii" family="Torpedinidae" genus="Torpedo" kingdom="Animalia" order="Torpediniformes" pageId="11" pageNumber="1199" phylum="Chordata" rank="genus">
<emphasis id="B935EA92227EB054208AED05FA8BFBAC" box="[1272,1362,1097,1120]" italics="true" pageId="11" pageNumber="1199">Torpedo</emphasis>
</taxonomicName>
and
<taxonomicName id="4C414D03227EB05421F7ED05FCABFB4C" authorityName="Dumeril" authorityYear="1852" class="Elasmobranchii" family="Hypnidae" genus="Hypnos" kingdom="Animalia" order="Torpediniformes" pageId="11" pageNumber="1199" phylum="Chordata" rank="genus">
<emphasis id="B935EA92227EB05421F7ED05FCABFB4C" italics="true" pageId="11" pageNumber="1199">Hypnos</emphasis>
</taxonomicName>
whereas in narcinoid genera the jaws are short and stout (
<bibRefCitation id="EFD04B71227EB05427EAEDC5FBECFB6C" author="Claeson, K. M." box="[920,1077,1161,1184]" pageId="11" pageNumber="1199" pagination="597 - 612" refId="ref20593" refString="Claeson, K. M. 2014. The impacts of comparative anatomy of electric rays (Batoidea: Torpediniformes) on their systematic hypotheses. Journal of Morphology, 275, 597 - 612." type="journal article" year="2014">Claeson 2014</bibRefCitation>
), the description of the jaws of †
<taxonomicName id="4C414D03227EB054272AEDE5FC03FB0C" authorityName="Carvalho" authorityYear="2010" box="[856,986,1193,1216]" class="Elasmobranchii" family="Narcinidae" genus="Titanonarke" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="11" pageNumber="1199" phylum="Chordata" rank="genus">
<emphasis id="B935EA92227EB054272AEDE5FC03FB0C" box="[856,986,1193,1216]" italics="true" pageId="11" pageNumber="1199">Titanonarke</emphasis>
</taxonomicName>
is further clarified here. The palatoquadrate of †
<taxonomicName id="4C414D03227EB0542709ED85FC24FB2C" authorityName="Carvalho" authorityYear="2010" box="[891,1021,1225,1248]" class="Elasmobranchii" family="Narcinidae" genus="Titanonarke" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="11" pageNumber="1199" phylum="Chordata" rank="genus">
<emphasis id="B935EA92227EB0542709ED85FC24FB2C" box="[891,1021,1225,1248]" italics="true" pageId="11" pageNumber="1199">Titanonarke</emphasis>
</taxonomicName>
is labiolingually compressed, narrower than the Meckels cartilage, and tapers towards the symphysis (
<figureCitation id="137A2A05227EB05427D3EC44FC38FAD3" box="[929,993,1288,1311]" captionStart="Figure 5" captionStartId="8.[159,227,682,703]" captionTargetBox="[365,1255,183,650]" captionTargetId="figure-236@8.[365,1261,183,650]" captionTargetPageId="8" captionText="Figure 5. †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site. A, MCSNV IG.VR.67290, close-up of the head and hyoid apparatus; B, reconstruction. Abbreviations: ao, antorbital cartilage; bb, basibranchials; bbc, basibranchial copula; cb, ceratobranchials; cc, chondrocranium; eb, epibranchials; hb, hypobranchials; hym, hyomandibula; la, labial cartilages; me, Meckels cartilage; nc, nasal capsule; pq, palatoquadrate; ps, pseudohyoid; ra, rostral appendix; rf, rostral fontanelle; ro, rostral cartilage; sca, scapulocoracoid; ss, suprascapula; syn, synarcual. Scale bars = 50 mm." figureDoi="http://doi.org/10.5281/zenodo.10883279" httpUri="https://zenodo.org/record/10883279/files/figure.png" pageId="11" pageNumber="1199">Fig. 5</figureCitation>
). Like the palatoquadrate of
<taxonomicName id="4C414D03227EB054216FEC44FC48FAF3" baseAuthorityName="Olfers" baseAuthorityYear="1831" class="Elasmobranchii" family="Narcinidae" genus="Narcine" kingdom="Animalia" order="Torpediniformes" pageId="11" pageNumber="1199" phylum="Chordata" rank="species" species="brasiliensis">
<emphasis id="B935EA92227EB054216FEC44FC48FAF3" italics="true" pageId="11" pageNumber="1199">Narcine brasiliensis</emphasis>
</taxonomicName>
(see
<bibRefCitation id="EFD04B71227EB05427BDEC64FB63FAF3" author="Dean, M. N. &amp; Motta, P. J." box="[975,1210,1320,1343]" pageId="11" pageNumber="1199" pagination="462 - 483" refId="ref21060" refString="Dean, M. N. &amp; Motta, P. J. 2004 a. Anatomy and functional morphology of the feeding apparatus of the lesser electric ray, Narcine brasiliensis (Elasmobranchii: Batoidea). Journal of Morphology, 262, 462 - 483." type="journal article" year="2004">Dean &amp; Motta 2004a</bibRefCitation>
), the palatoquadrate of †
<taxonomicName id="4C414D03227EB054272AEC04FC60FA93" baseAuthorityName="Jaekel" baseAuthorityYear="1894" box="[856,953,1352,1375]" class="Elasmobranchii" family="Narcinidae" genus="Titanonarke" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="11" pageNumber="1199" phylum="Chordata" rank="species" species="molini">
<emphasis id="B935EA92227EB054272AEC04FC60FA93" box="[856,953,1352,1375]" italics="true" pageId="11" pageNumber="1199">T. molini</emphasis>
</taxonomicName>
possesses a strong condyle that articulates with the Meckels cartilage at the mandibular articular fossa
<emphasis id="B935EA92227EB054273FECC4FC51FA53" box="[845,904,1416,1439]" italics="true" pageId="11" pageNumber="1199">sensu</emphasis>
<bibRefCitation id="EFD04B71227EB05427FDECC4FB5FFA53" author="Dean, M. N. &amp; Motta, P. J." box="[911,1158,1416,1439]" pageId="11" pageNumber="1199" pagination="462 - 483" refId="ref21060" refString="Dean, M. N. &amp; Motta, P. J. 2004 a. Anatomy and functional morphology of the feeding apparatus of the lesser electric ray, Narcine brasiliensis (Elasmobranchii: Batoidea). Journal of Morphology, 262, 462 - 483." type="journal article" year="2004">Dean &amp; Motta (2004a)</bibRefCitation>
. The Meckels cartilages are stout, flat and broad. There are two pairs of small, slender and subtriangular labial cartilages situated near the symphysis of the jaws, surrounding the tooth bands (
<figureCitation id="137A2A05227EB0542118ECA4FA72FA33" box="[1386,1451,1512,1535]" captionStart="Figure 5" captionStartId="8.[159,227,682,703]" captionTargetBox="[365,1255,183,650]" captionTargetId="figure-236@8.[365,1261,183,650]" captionTargetPageId="8" captionText="Figure 5. †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site. A, MCSNV IG.VR.67290, close-up of the head and hyoid apparatus; B, reconstruction. Abbreviations: ao, antorbital cartilage; bb, basibranchials; bbc, basibranchial copula; cb, ceratobranchials; cc, chondrocranium; eb, epibranchials; hb, hypobranchials; hym, hyomandibula; la, labial cartilages; me, Meckels cartilage; nc, nasal capsule; pq, palatoquadrate; ps, pseudohyoid; ra, rostral appendix; rf, rostral fontanelle; ro, rostral cartilage; sca, scapulocoracoid; ss, suprascapula; syn, synarcual. Scale bars = 50 mm." figureDoi="http://doi.org/10.5281/zenodo.10883279" httpUri="https://zenodo.org/record/10883279/files/figure.png" pageId="11" pageNumber="1199">Fig. 5</figureCitation>
). Combined, upper and lower labial cartilages are less than the length of the Meckels cartilage. The triangular element displaced to the corner of the left jaw joint in
<collectionCode id="ED50AE45227EB054273FEF2BFC72F9B2" box="[845,939,1639,1662]" pageId="11" pageNumber="1199">MCSNV</collectionCode>
<collectionCode id="ED50AE45227EB05427C7EF2BFC0AF9B2" box="[949,979,1639,1662]" country="China" name="Institute of Geology" pageId="11" pageNumber="1199">IG</collectionCode>
.135576, and interpreted by
<bibRefCitation id="EFD04B71227EB0542178EF2BFA6DF9B2" author="Carvalho, M. R. de" box="[1290,1460,1639,1662]" pageId="11" pageNumber="1199" refId="ref19921" refString="Carvalho, M. R. de. 2010. Morphology and phylogenetic relationships of the giant electric ray from the Eocene of Monte Bolca, Italy (Chondrichthyes: Torpediniformes). Pp. 183 - 198 in D. K. Elliot, J. G. Maisey, X. Yu &amp; D. Miao (eds) Morphology, phylogeny and paleobiogeography of fossil fishes. Verlag Dr. Friedrich Pfeil, Munich." type="book" year="2010">Carvalho (2010</bibRefCitation>
, fig. 5b) as a labial cartilage, instead appears to be a dorsal flange of the sustentanculum of the Meckels cartilage. Both jaws are not fused medially.
</paragraph>
<caption id="DF3E6608227EB05424EDEFB8FF30F842" ID-DOI="http://doi.org/10.5281/zenodo.10883299" ID-Zenodo-Dep="10883299" httpUri="https://zenodo.org/record/10883299/files/figure.png" pageId="11" pageNumber="1199" startId="11.[159,227,1780,1801]" targetBox="[166,774,1083,1748]" targetPageId="11" targetType="figure">
<paragraph id="8BFE3680227EB05424EDEFB8FF30F842" blockId="11.[159,781,1779,1934]" pageId="11" pageNumber="1199">
<emphasis id="B935EA92227EB05424EDEFB8FED1F8C5" bold="true" box="[159,264,1780,1801]" pageId="11" pageNumber="1199">Figure 12.</emphasis>
<taxonomicName id="4C414D03227EB054256CEFB8FDBFF8C5" authority="(Jaekel, 1894)" baseAuthorityName="Jaekel" baseAuthorityYear="1894" box="[286,614,1780,1801]" class="Elasmobranchii" family="Narcinidae" genus="Titanonarke" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="11" pageNumber="1199" phylum="Chordata" rank="species" species="molini">
<emphasis id="B935EA92227EB054256CEFB8FE0EF8C5" box="[286,471,1780,1801]" italics="true" pageId="11" pageNumber="1199">Titanonarke molini</emphasis>
(
<bibRefCitation id="EFD04B71227EB0542596EFB8FD87F8C5" author="Jaekel, O." box="[484,606,1780,1801]" pageId="11" pageNumber="1199" refId="ref23357" refString="Jaekel, O. 1894. Die eoca ¨ nen Selachier vom Monte Bolca: ein Beitrag zur Morphogenie der Wirbelthiere. Springer, Berlin, 177 pp." type="book" year="1894">Jaekel, 1894</bibRefCitation>
)
</taxonomicName>
from the Eocene Monte Postale site. Close-up of the distal end of pelvic fins of MCSNV IG.91128, which is supposed to be the unique male individual based on the presence of claspers. Abbreviations: cla, clasper; dfr, first dorsal-fin radials, pelvic radials. Scale bar = 50 mm.
</paragraph>
</caption>
<paragraph id="8BFE3680227EB0532707EFBBFA8DFC1E" blockId="11.[845,1467,1783,1934]" lastBlockId="12.[845,1467,923,1615]" lastPageId="12" lastPageNumber="1200" pageId="11" pageNumber="1199">
<emphasis id="B935EA92227EB0542707EFBBFBA8F8C2" bold="true" box="[885,1137,1783,1806]" pageId="11" pageNumber="1199">Hyoid and gill arches.</emphasis>
The exquisite preservation of
<collectionCode id="ED50AE45227EB054273FEE5BFC72F8E2" box="[845,939,1815,1838]" pageId="11" pageNumber="1199">MCSNV</collectionCode>
<collectionCode id="ED50AE45227EB05427CAEE5BFC0CF8E2" box="[952,981,1815,1838]" country="China" name="Institute of Geology" pageId="11" pageNumber="1199">IG</collectionCode>
.VR.67290 allows a detailed description of most of the hyoid arch (
<figureCitation id="137A2A05227EB054202FEE7BFB47F882" box="[1117,1182,1847,1870]" captionStart="Figure 5" captionStartId="8.[159,227,682,703]" captionTargetBox="[365,1255,183,650]" captionTargetId="figure-236@8.[365,1261,183,650]" captionTargetPageId="8" captionText="Figure 5. †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site. A, MCSNV IG.VR.67290, close-up of the head and hyoid apparatus; B, reconstruction. Abbreviations: ao, antorbital cartilage; bb, basibranchials; bbc, basibranchial copula; cb, ceratobranchials; cc, chondrocranium; eb, epibranchials; hb, hypobranchials; hym, hyomandibula; la, labial cartilages; me, Meckels cartilage; nc, nasal capsule; pq, palatoquadrate; ps, pseudohyoid; ra, rostral appendix; rf, rostral fontanelle; ro, rostral cartilage; sca, scapulocoracoid; ss, suprascapula; syn, synarcual. Scale bars = 50 mm." figureDoi="http://doi.org/10.5281/zenodo.10883279" httpUri="https://zenodo.org/record/10883279/files/figure.png" pageId="11" pageNumber="1199">Fig. 5</figureCitation>
). The hyomandibulae are narrow and elongate, slightly stout at the proximal base, and tapering distally towards their articulation with the Meckels cartilage. The dorsal and ventral pseudohyoids are long and slender, and located just posterior to the articulation of the hyomandibula with the otic region of the chondrocranium. †
<taxonomicName id="4C414D032279B05325F5EAB6FDD0FBDD" authorityName="Carvalho" authorityYear="2010" box="[391,521,1018,1041]" class="Elasmobranchii" family="Narcinidae" genus="Titanonarke" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="12" pageNumber="1200" phylum="Chordata" rank="genus">
<emphasis id="B935EA922279B05325F5EAB6FDD0FBDD" box="[391,521,1018,1041]" italics="true" pageId="12" pageNumber="1200">Titanonarke</emphasis>
</taxonomicName>
lacks ceratohyals, resembling the condition of
<taxonomicName id="4C414D032279B05320DBEAD7FB26FC7E" baseAuthorityName="Claeson" baseAuthorityYear="2014" box="[1193,1279,923,946]" class="Elasmobranchii" family="Narcinidae" genus="Narcine" kingdom="Animalia" order="Torpediniformes" pageId="12" pageNumber="1200" phylum="Chordata" rank="genus">
<emphasis id="B935EA922279B05320DBEAD7FB26FC7E" box="[1193,1279,923,946]" italics="true" pageId="12" pageNumber="1200">Narcine</emphasis>
</taxonomicName>
and
<taxonomicName id="4C414D032279B0532138EAD7FA63FC7E" authorityName="Heckel" authorityYear="1846" box="[1354,1466,923,946]" class="Elasmobranchii" family="Narcinidae" genus="Discopyge" kingdom="Animalia" order="Torpediniformes" pageId="12" pageNumber="1200" phylum="Chordata" rank="genus">
<emphasis id="B935EA922279B0532138EAD7FA63FC7E" box="[1354,1466,923,946]" italics="true" pageId="12" pageNumber="1200">Discopyge</emphasis>
</taxonomicName>
among narcinoids (Miyake &amp; McEachran 1991).
</paragraph>
<caption id="DF3E66082279B05324EDEBB5FE4CFC92" ID-DOI="http://doi.org/10.5281/zenodo.10883301" ID-Zenodo-Dep="10883301" httpUri="https://zenodo.org/record/10883301/files/figure.png" pageId="12" pageNumber="1200" startId="12.[159,227,761,782]" targetBox="[317,1309,183,729]" targetPageId="12" targetType="figure">
<paragraph id="8BFE36802279B05324EDEBB5FE4CFC92" blockId="12.[159,1467,760,862]" pageId="12" pageNumber="1200">
<emphasis id="B935EA922279B05324EDEBB5FED3FCC2" bold="true" box="[159,266,761,782]" pageId="12" pageNumber="1200">Figure 13.</emphasis>
<taxonomicName id="4C414D032279B0532552EBB5FDB4FCC2" authority="(Jaekel, 1894)" baseAuthorityName="Jaekel" baseAuthorityYear="1894" box="[288,621,761,782]" class="Elasmobranchii" family="Narcinidae" genus="Titanonarke" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="12" pageNumber="1200" phylum="Chordata" rank="species" species="molini">
<emphasis id="B935EA922279B0532552EBB5FE03FCC2" box="[288,474,761,782]" italics="true" pageId="12" pageNumber="1200">Titanonarke molini</emphasis>
(
<bibRefCitation id="EFD04B712279B053259BEBB5FDBCFCC2" author="Jaekel, O." box="[489,613,761,782]" pageId="12" pageNumber="1200" refId="ref23357" refString="Jaekel, O. 1894. Die eoca ¨ nen Selachier vom Monte Bolca: ein Beitrag zur Morphogenie der Wirbelthiere. Springer, Berlin, 177 pp." type="book" year="1894">Jaekel, 1894</bibRefCitation>
)
</taxonomicName>
from the Eocene Monte Postale site. Details of the precaudal tail of
<emphasis id="B935EA922279B053216DEBB5FAEFFCC2" bold="true" box="[1311,1334,761,782]" pageId="12" pageNumber="1200">A,</emphasis>
MCSNV IG. VR.67290 and
<emphasis id="B935EA922279B0532540EA58FE91FCE5" bold="true" box="[306,328,788,809]" pageId="12" pageNumber="1200">B,</emphasis>
MGP-PD 26276. The arrows mark the position of the two dorsal fins in MCSNV IG.VR.67290 and the second dorsal fin in MGP-PD 26276 already detected and figured by
<bibRefCitation id="EFD04B712279B05326CBEA62FCE7FC8F" author="Jaekel, O." box="[697,830,814,835]" pageId="12" pageNumber="1200" refId="ref23357" refString="Jaekel, O. 1894. Die eoca ¨ nen Selachier vom Monte Bolca: ein Beitrag zur Morphogenie der Wirbelthiere. Springer, Berlin, 177 pp." type="book" year="1894">Jaekel (1894)</bibRefCitation>
.
<emphasis id="B935EA922279B0532738EA62FCB8FC8F" bold="true" box="[842,865,814,835]" pageId="12" pageNumber="1200">C,</emphasis>
Detail of the caudal fin of MCSNV IG.91129. Scale bars: A, B = 50 mm; C = 10 mm.
</paragraph>
</caption>
<paragraph id="8BFE36802279B0532715EA96FA9FF983" blockId="12.[845,1467,923,1615]" pageId="12" pageNumber="1200">
There are five pairs of ceratobranchials. The anterior four pairs are large, and have a central depression (or fossa) with a small fenestra for the insertion of the depressor muscles (see Carvalho &amp; Śeret 2002). The fenestrae of †
<taxonomicName id="4C414D032279B053272AED16FC03FBBD" authorityName="Carvalho" authorityYear="2010" box="[856,986,1114,1137]" class="Elasmobranchii" family="Narcinidae" genus="Titanonarke" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="12" pageNumber="1200" phylum="Chordata" rank="genus">
<emphasis id="B935EA922279B053272AED16FC03FBBD" box="[856,986,1114,1137]" italics="true" pageId="12" pageNumber="1200">Titanonarke</emphasis>
</taxonomicName>
appear larger in small individuals (
<figureCitation id="137A2A052279B0532118ED16FA72FBBD" box="[1386,1451,1114,1137]" captionStart="Figure 6" captionStartId="8.[159,227,1806,1827]" captionTargetBox="[365,1252,1347,1774]" captionTargetId="figure-317@8.[365,1261,1347,1774]" captionTargetPageId="8" captionText="Figure 6. †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site. A, the juvenile individual MCSNV IG.VR.91359; B, reconstruction. Abbreviations: ao, antorbital cartilage; cb, ceratobranchials; hym, hyomandibula; la, labial cartilages; me, Meckels cartilage; mes, mesopterygium; met, metapterygium; nc, nasal capsule; pel, prepelvic process; pub, puboischiadic bar; pq, palatoquadrate; pro, propterygium; ps, pseudohyoid; rad, pectoral radials; ro, rostral cartilage; sca, scapulocoracoid; ss, suprascapula; syn, synarcual. Scale bars = 50 mm." figureDoi="http://doi.org/10.5281/zenodo.10883285" httpUri="https://zenodo.org/record/10883285/files/figure.png" pageId="12" pageNumber="1200">Fig. 6</figureCitation>
), as in
<taxonomicName id="4C414D032279B05327E2ED36FBD8FB5D" baseAuthorityName="Fechhelm &amp; McEachran" baseAuthorityYear="1984" box="[912,1025,1146,1169]" class="Elasmobranchii" family="Narcinidae" genus="Diplobatis" kingdom="Animalia" order="Torpediniformes" pageId="12" pageNumber="1200" phylum="Chordata" rank="genus">
<emphasis id="B935EA922279B05327E2ED36FBD8FB5D" box="[912,1025,1146,1169]" italics="true" pageId="12" pageNumber="1200">Diplobatis</emphasis>
</taxonomicName>
(Miyake &amp; McEachran 1991, fig. 6H;
<bibRefCitation id="EFD04B712279B053273FEDD6FC39FB7D" author="Claeson, K. M." box="[845,992,1178,1201]" pageId="12" pageNumber="1200" pagination="597 - 612" refId="ref20593" refString="Claeson, K. M. 2014. The impacts of comparative anatomy of electric rays (Batoidea: Torpediniformes) on their systematic hypotheses. Journal of Morphology, 275, 597 - 612." type="journal article" year="2014">Claeson 2014</bibRefCitation>
, supplemental pl. 12). The fifth ceratobranchials are slender and posteriorly oriented, and articulate with the scapular process of the scapulocoracoid. The epibranchial elements are not clearly recognizable, being partially covered by the ceratohyals. At least three small basibranchials are recognizable. The basibranchial copula is large and rounded, with a small caudal tip or tab in its posterior margin. We counted five pairs of ovoid or beanlike hypobranchials. Hypobranchials are segmented, resembling the plesiomorphic condition of
<taxonomicName id="4C414D032279B0532146ECF5FA6AFA1C" authorityName="Alcock" authorityYear="1898" box="[1332,1459,1465,1488]" class="Elasmobranchii" family="Narcinidae" genus="Benthobatis" kingdom="Animalia" order="Torpediniformes" pageId="12" pageNumber="1200" phylum="Chordata" rank="genus">
<emphasis id="B935EA922279B0532146ECF5FA6AFA1C" box="[1332,1459,1465,1488]" italics="true" pageId="12" pageNumber="1200">Benthobatis</emphasis>
</taxonomicName>
,
<taxonomicName id="4C414D032279B053273FEC94FC64FA23" authorityName="Heckel" authorityYear="1846" box="[845,957,1496,1519]" class="Elasmobranchii" family="Narcinidae" genus="Discopyge" kingdom="Animalia" order="Torpediniformes" pageId="12" pageNumber="1200" phylum="Chordata" rank="genus">
<emphasis id="B935EA922279B053273FEC94FC64FA23" box="[845,957,1496,1519]" italics="true" pageId="12" pageNumber="1200">Discopyge</emphasis>
</taxonomicName>
and
<taxonomicName id="4C414D032279B0532780EC94FBA0FA23" authorityName="Regan" authorityYear="1921" box="[1010,1145,1496,1519]" class="Elasmobranchii" family="Narkidae" genus="Heteronarce" kingdom="Animalia" order="Torpediniformes" pageId="12" pageNumber="1200" phylum="Chordata" rank="genus">
<emphasis id="B935EA922279B0532780EC94FBA0FA23" box="[1010,1145,1496,1519]" italics="true" pageId="12" pageNumber="1200">Heteronarce</emphasis>
</taxonomicName>
among narcinoids (Miyake &amp; McEachran 1991, fig. 6F, G;
<bibRefCitation id="EFD04B712279B05320F9ECB4FAC4F9C3" author="Claeson, K. M." box="[1163,1309,1528,1551]" pageId="12" pageNumber="1200" pagination="597 - 612" refId="ref20593" refString="Claeson, K. M. 2014. The impacts of comparative anatomy of electric rays (Batoidea: Torpediniformes) on their systematic hypotheses. Journal of Morphology, 275, 597 - 612." type="journal article" year="2014">Claeson 2014</bibRefCitation>
, supplemental pl. 12). Pharyngobranchials, extrabranchials and branchial rays are not preserved in the available material.
</paragraph>
<paragraph id="8BFE36802279B0522707EF34FF2DF942" blockId="12.[845,1467,1656,1934]" lastBlockId="13.[159,781,1368,1933]" lastPageId="13" lastPageNumber="1201" pageId="12" pageNumber="1200">
<emphasis id="B935EA922279B0532707EF34FAD2F943" bold="true" box="[885,1291,1656,1679]" pageId="12" pageNumber="1200">Synarcual and vertebral column.</emphasis>
The synarcual cartilage is strongly calcified in all mature specimens, though not in the embryo preserved inside the abdominal cavity of the
<typeStatus id="54FA88222279B0532782EF94FB94F923" box="[1008,1101,1752,1775]" pageId="12" pageNumber="1200" type="holotype">holotype</typeStatus>
(see the section Embryo). In mature specimens the synarcual exhibits the typical prismatic tessellated cartilage found in elasmobranchs. The posteroventralmost portion of tessellated cartilage flanks several mineralized vertebral centra, which comprise densely packed areolar cartilage. Anteriorly, the synarcual cartilage contacts the occipital condyles of the chondrocranium via the occipital cotyles. The morphology of the anterior portion of the synarcual is difficult to discern, it being partially hidden by the hyoid arch (most of the specimens are ventrally exposed). Therefore, the position of the dorsal rim of the anterior neural canal opening (synarcual mouth) with respect to the occipital cotyle and the lips, as well as the shape of the ventral rim of the anterior neural canal opening (synarcual lip), are difficult to discern.
</paragraph>
<caption id="DF3E66082279B05324EDEE42FD30F842" ID-DOI="http://doi.org/10.5281/zenodo.10883305" ID-Zenodo-Dep="10883305" httpUri="https://zenodo.org/record/10883305/files/figure.png" pageId="12" pageNumber="1200" startId="12.[159,227,1806,1827]" targetBox="[166,774,1108,1775]" targetPageId="12" targetType="figure">
<paragraph id="8BFE36802279B05324EDEE42FD30F842" blockId="12.[159,781,1806,1934]" pageId="12" pageNumber="1200">
<emphasis id="B935EA922279B05324EDEE42FED1F8EF" bold="true" box="[159,264,1806,1827]" pageId="12" pageNumber="1200">Figure 14.</emphasis>
<taxonomicName id="4C414D032279B053256CEE42FDBFF8EF" authority="(Jaekel, 1894)" baseAuthorityName="Jaekel" baseAuthorityYear="1894" box="[286,614,1806,1827]" class="Elasmobranchii" family="Narcinidae" genus="Titanonarke" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="12" pageNumber="1200" phylum="Chordata" rank="species" species="molini">
<emphasis id="B935EA922279B053256CEE42FE0EF8EF" box="[286,471,1806,1827]" italics="true" pageId="12" pageNumber="1200">Titanonarke molini</emphasis>
(
<bibRefCitation id="EFD04B712279B0532596EE42FD87F8EF" author="Jaekel, O." box="[484,606,1806,1827]" pageId="12" pageNumber="1200" refId="ref23357" refString="Jaekel, O. 1894. Die eoca ¨ nen Selachier vom Monte Bolca: ein Beitrag zur Morphogenie der Wirbelthiere. Springer, Berlin, 177 pp." type="book" year="1894">Jaekel, 1894</bibRefCitation>
)
</taxonomicName>
from the Eocene of Monte Postale site.
<emphasis id="B935EA922279B05325E4EE65FE77F8F2" bold="true" box="[406,430,1833,1854]" pageId="12" pageNumber="1200">A,</emphasis>
upper and lower tooth bands in MCSNV IG.VR.67290, with a close-up of some teeth in the area indicated.
<emphasis id="B935EA922279B0532574EE12FEC5F8BF" bold="true" box="[262,284,1886,1907]" pageId="12" pageNumber="1200">B,</emphasis>
reconstruction. Abbreviations: la, labial cartilage; me, Meckels cartilage; pq, palatoquadrate. Scale bars 5 mm.
</paragraph>
</caption>
<caption id="DF3E66082278B05224EDEDC6FDE8FAE8" ID-DOI="http://doi.org/10.5281/zenodo.10883307" ID-Zenodo-Dep="10883307" httpUri="https://zenodo.org/record/10883307/files/figure.png" pageId="13" pageNumber="1201" startId="13.[159,227,1162,1183]" targetBox="[173,1451,183,1130]" targetPageId="13" targetType="figure">
<paragraph id="8BFE36802278B05224EDEDC6FDE8FAE8" blockId="13.[159,1468,1161,1316]" pageId="13" pageNumber="1201">
<emphasis id="B935EA922278B05224EDEDC6FE9CFB53" bold="true" box="[159,325,1162,1183]" pageId="13" pageNumber="1201">Figure 15. A, B,</emphasis>
<taxonomicName id="4C414D032278B0522524EDC6FD45FB53" authority="(Jaekel, 1894)" baseAuthorityName="Jaekel" baseAuthorityYear="1894" box="[342,668,1162,1183]" class="Elasmobranchii" family="Narcinidae" genus="Titanonarke" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="13" pageNumber="1201" phylum="Chordata" rank="species" species="molini">
<emphasis id="B935EA922278B0522524EDC6FDD7FB53" box="[342,526,1162,1183]" italics="true" pageId="13" pageNumber="1201">Titanonarke molini</emphasis>
(
<bibRefCitation id="EFD04B712278B0522669EDC6FD4DFB53" author="Jaekel, O." box="[539,660,1162,1183]" pageId="13" pageNumber="1201" refId="ref23357" refString="Jaekel, O. 1894. Die eoca ¨ nen Selachier vom Monte Bolca: ein Beitrag zur Morphogenie der Wirbelthiere. Springer, Berlin, 177 pp." type="book" year="1894">Jaekel, 1894</bibRefCitation>
)
</taxonomicName>
from the Eocene Monte Postale site;
<emphasis id="B935EA922278B0522076EDC6FBC2FB53" bold="true" box="[1028,1051,1162,1183]" pageId="13" pageNumber="1201">A,</emphasis>
detail of the abdominal region in MGP-PD 26275 showing the stomach content;
<emphasis id="B935EA922278B0522679EDE8FDF8FB75" bold="true" box="[523,545,1188,1209]" pageId="13" pageNumber="1201">B,</emphasis>
reconstruction; note also the embryo lying next to the stomach.
<emphasis id="B935EA922278B05220EAEDE8FB14FB75" bold="true" box="[1176,1229,1188,1209]" pageId="13" pageNumber="1201">C, D,</emphasis>
close-up of some of the larger foraminifera of the genus †
<taxonomicName id="4C414D032278B052259AEDF3FD9DFB18" authorityName="d'Orbigny" authorityYear="1826" box="[488,580,1215,1236]" class="Magnoliopsida" family="Loranthaceae" genus="Alveolina" kingdom="Plantae" order="Santalales" pageId="13" pageNumber="1201" phylum="Tracheophyta" rank="genus">
<emphasis id="B935EA922278B052259AEDF3FD9DFB18" box="[488,580,1215,1236]" italics="true" pageId="13" pageNumber="1201">Alveolina</emphasis>
</taxonomicName>
in the stomach of MGP-PD 26275.
<emphasis id="B935EA922278B05227DAEDF3FC67FB18" bold="true" box="[936,958,1215,1236]" pageId="13" pageNumber="1201">E,</emphasis>
dissected specimen of
<taxonomicName id="4C414D032278B05220D0EDF3FA8FFB18" authorityName="Bonaparte" authorityYear="1835" box="[1186,1366,1215,1236]" class="Elasmobranchii" family="Torpedinidae" genus="Torpedo" kingdom="Animalia" order="Torpediniformes" pageId="13" pageNumber="1201" phylum="Chordata" rank="species" species="nobiliana">
<emphasis id="B935EA922278B05220D0EDF3FA8FFB18" box="[1186,1366,1215,1236]" italics="true" pageId="13" pageNumber="1201">Torpedo nobiliana</emphasis>
</taxonomicName>
in ventral view (ESB tn200707_159) showing the position of the stomach, used to identify the accumulation in MGP-PD 26275 as gut contents. Abbreviations: int, intestine; liv, liver; met, metapterygium; pub, puboischiadic bar; sca, scapulocoracoid; st, stomach, syn synarcual. Scale bars: A, B = 50 mm; C, D = 2 mm.
</paragraph>
</caption>
<paragraph id="8BFE36802278B05224C8EFDBFBB5FA03" blockId="13.[159,781,1368,1933]" lastBlockId="13.[845,1467,1368,1933]" pageId="13" pageNumber="1201">
The synarcual possesses lateral stays, which are posteriorly displaced (
<figureCitation id="137A2A052278B0522535EFFBFE5CF902" box="[327,389,1719,1742]" captionStart="Figure 9" captionStartId="10.[159,227,702,723]" captionTargetBox="[365,1261,183,670]" captionTargetId="figure-180@10.[365,1261,183,670]" captionTargetPageId="10" captionText="Figure 9. A, synarcual and pectoral girdle of Narcine brasiliensis (AMNH 95343) in dorsal view; B, synarcual and pectoral girdle of †Titanonarke molini (MCSNV IG.VR.91359) in ventral view. The arrowheads indicate the posteriorly directed lateral stays of the synarcual. The arrows indicate the posteriorly directed scapular process of the scapulocoracoid. Abbreviations: cb, ceratocranchials; sca, scapulocoracoid; ss, suprascapula; syn, synarcual. Scale bar = 5 mm." figureDoi="http://doi.org/10.5281/zenodo.10883293" httpUri="https://zenodo.org/record/10883293/files/figure.png" pageId="13" pageNumber="1201">Fig. 9</figureCitation>
) as in
<taxonomicName id="4C414D032278B05225BDEFFBFD97F902" authorityName="Alcock" authorityYear="1898" box="[463,590,1719,1742]" class="Elasmobranchii" family="Narcinidae" genus="Benthobatis" kingdom="Animalia" order="Torpediniformes" pageId="13" pageNumber="1201" phylum="Chordata" rank="genus">
<emphasis id="B935EA922278B05225BDEFFBFD97F902" box="[463,590,1719,1742]" italics="true" pageId="13" pageNumber="1201">Benthobatis</emphasis>
</taxonomicName>
,
<taxonomicName id="4C414D032278B052262EEFFBFD15F902" authorityName="Heckel" authorityYear="1846" box="[604,716,1719,1742]" class="Elasmobranchii" family="Narcinidae" genus="Discopyge" kingdom="Animalia" order="Torpediniformes" pageId="13" pageNumber="1201" phylum="Chordata" rank="genus">
<emphasis id="B935EA922278B052262EEFFBFD15F902" box="[604,716,1719,1742]" italics="true" pageId="13" pageNumber="1201">Discopyge</emphasis>
</taxonomicName>
,
<taxonomicName id="4C414D032278B05226A9EFFBFF12F921" baseAuthorityName="Claeson" baseAuthorityYear="2014" class="Elasmobranchii" family="Narcinidae" genus="Narcine" kingdom="Animalia" order="Torpediniformes" pageId="13" pageNumber="1201" phylum="Chordata" rank="genus">
<emphasis id="B935EA922278B05226A9EFFBFF12F921" italics="true" pageId="13" pageNumber="1201">Narcine</emphasis>
</taxonomicName>
,
<taxonomicName id="4C414D032278B05224AFEF9AFEBDF921" authorityName="Regan" authorityYear="1921" box="[221,356,1750,1773]" class="Elasmobranchii" family="Narkidae" genus="Heteronarce" kingdom="Animalia" order="Torpediniformes" pageId="13" pageNumber="1201" phylum="Chordata" rank="genus">
<emphasis id="B935EA922278B05224AFEF9AFEBDF921" box="[221,356,1750,1773]" italics="true" pageId="13" pageNumber="1201">Heteronarce</emphasis>
</taxonomicName>
and
<taxonomicName id="4C414D032278B05225EFEF9AFD48F921" authority="(Claeson 2014)" baseAuthorityName="Claeson" baseAuthorityYear="2014" box="[413,657,1750,1773]" class="Elasmobranchii" family="Narkidae" genus="Narke" kingdom="Animalia" order="Torpediniformes" pageId="13" pageNumber="1201" phylum="Chordata" rank="genus">
<emphasis id="B935EA922278B05225EFEF9AFE07F921" box="[413,478,1750,1773]" italics="true" pageId="13" pageNumber="1201">Narke</emphasis>
(
<bibRefCitation id="EFD04B712278B0522582EF9AFD5FF921" author="Claeson, K. M." box="[496,646,1750,1773]" pageId="13" pageNumber="1201" pagination="597 - 612" refId="ref20593" refString="Claeson, K. M. 2014. The impacts of comparative anatomy of electric rays (Batoidea: Torpediniformes) on their systematic hypotheses. Journal of Morphology, 275, 597 - 612." type="journal article" year="2014">Claeson 2014</bibRefCitation>
)
</taxonomicName>
. The distal end of the lateral stays is apparently tab-like and their anterior margin forms an obtuse angle with the anterolateral margin of the synarcual. The long posterior flanges of the synarcual surround the anteriormost 812 free vertebral centra, and their posterior margins end posteriorly to the scapulocoracoid bar, resembling the condition of nonnarkid torpediniforms (
<bibRefCitation id="EFD04B712278B0522036EC34FB2FFA43" author="Compagno, L. J. V." box="[1092,1270,1400,1423]" pageId="13" pageNumber="1201" refId="ref20733" refString="Compagno, L. J. V. 1999. Chapter 3. Endoskeleton. Pp. 69 - 92 in W. C. Hamlett (ed.) Sharks, skates and rays. The biology of elasmobranch fishes. Johns Hopkins Press, Baltimore." type="book" year="1999">Compagno 1999</bibRefCitation>
). It is not possible to detect the number of fused vertebrae that constitute the synarcual, or the foramina.
</paragraph>
<paragraph id="8BFE36802278B0522715EC9BFB49F8E1" blockId="13.[845,1467,1368,1933]" pageId="13" pageNumber="1201">
The vertebral column of †
<taxonomicName id="4C414D032278B05220FDEC9BFB2AFA22" baseAuthorityName="Jaekel" baseAuthorityYear="1894" box="[1167,1267,1495,1518]" class="Elasmobranchii" family="Narcinidae" genus="Titanonarke" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="13" pageNumber="1201" phylum="Chordata" rank="species" species="molini">
<emphasis id="B935EA922278B05220FDEC9BFB2AFA22" box="[1167,1267,1495,1518]" italics="true" pageId="13" pageNumber="1201">T. molini</emphasis>
</taxonomicName>
consists of about 153155 vertebral centra; of these, 2730 are trunk centra (1820% of total, from the first distinguishable centrum to the anterior margin of the puboischiadic bar), 100115 are precaudals (6575%, from the anterior margin of the puboischiadic bars to the upper origin of the caudal fin), and 2325 are caudals (1516%, from the upper caudal fin origin to the end of the series). The number of vertebrae is by far the largest compared to all living torpediniforms and can be considered an autapomorphic condition of †
<taxonomicName id="4C414D032278B0522707EE5AFC2EF8E1" authorityName="Carvalho" authorityYear="2010" box="[885,1015,1814,1837]" class="Elasmobranchii" family="Narcinidae" genus="Titanonarke" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="13" pageNumber="1201" phylum="Chordata" rank="genus">
<emphasis id="B935EA922278B0522707EE5AFC2EF8E1" box="[885,1015,1814,1837]" italics="true" pageId="13" pageNumber="1201">Titanonarke</emphasis>
</taxonomicName>
(see
<tableCitation id="C6C3033B2278B052205DEE5AFBA6F8E1" box="[1071,1151,1814,1837]" captionStart="Table 2" captionStartId="16.[159,217,186,207]" captionTargetPageId="16" captionText="Table 2. Summary of selected meristic features used to discriminate fossil and living genera of the family Narcinidae. Includes new data from the examined material and data from Fechhelm &amp; McEachran (1984), Rincon (1997), Carvalho et al. (1999, 2002a, b, 2003), Carvalho (2001, 2008), Rincon et al. (2001), Carvalho &amp; Śeret (2002), Carvalho &amp; Randall (2003), Menni et al. (2008), Carvalho &amp; White (2016), Last et al. (2016), Froese &amp; Pauly (2017)." pageId="13" pageNumber="1201">Table 2</tableCitation>
).
</paragraph>
<paragraph id="8BFE36802278B0512715EE7AFD9CFA7C" blockId="13.[845,1467,1368,1933]" lastBlockId="14.[159,781,1241,1456]" lastPageId="14" lastPageNumber="1202" pageId="13" pageNumber="1201">
The vertebral centra are strongly calcified, subrectangular in shape and anteroposteriorly short. Large basiventral processes are visible along most of the vertebral centra, from the posterior tip of the synarcual to the caudal fin base. There are eight to 10 pairs of ribs articulating with centra posterior to the puboischiadic bar (
<figureCitation id="137A2A05227BB0512618EC55FD16FAFC" box="[618,719,1305,1328]" captionStart="Figure 11" captionStartId="11.[159,227,701,722]" captionTargetBox="[365,1257,183,669]" captionTargetId="figure-384@11.[365,1261,183,669]" captionTargetPageId="11" captionText="Figure 11. Pelvic fins of †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site. A, MCSNV IG.VR.67290; B, reconstruction. Abbreviations: bas, basipterygia; ilp, iliac process; pel, pelvic processes; pub, puboischiadic bar; rad, pelvic radials. The arrowhead indicates the iliac process. Scale bars = 50 mm." figureDoi="http://doi.org/10.5281/zenodo.10883297" httpUri="https://zenodo.org/record/10883297/files/figure.png" pageId="14" pageNumber="1202">Fig. 11A</figureCitation>
). The low number of rib pairs is similar to that of
<taxonomicName id="4C414D03227BB05126F5EC75FCDFFA9C" authorityName="Alcock" authorityYear="1898" box="[647,774,1337,1360]" class="Elasmobranchii" family="Narcinidae" genus="Benthobatis" kingdom="Animalia" order="Torpediniformes" pageId="14" pageNumber="1202" phylum="Chordata" rank="genus">
<emphasis id="B935EA92227BB05126F5EC75FCDFFA9C" box="[647,774,1337,1360]" italics="true" pageId="14" pageNumber="1202">Benthobatis</emphasis>
</taxonomicName>
,
<taxonomicName id="4C414D03227BB05124EDEC15FED6FABC" authorityName="Heckel" authorityYear="1846" box="[159,271,1369,1392]" class="Elasmobranchii" family="Narcinidae" genus="Discopyge" kingdom="Animalia" order="Torpediniformes" pageId="14" pageNumber="1202" phylum="Chordata" rank="genus">
<emphasis id="B935EA92227BB05124EDEC15FED6FABC" box="[159,271,1369,1392]" italics="true" pageId="14" pageNumber="1202">Discopyge</emphasis>
</taxonomicName>
,
<taxonomicName id="4C414D03227BB0512550EC15FE4AFABC" baseAuthorityName="Fechhelm &amp; McEachran" baseAuthorityYear="1984" box="[290,403,1369,1392]" class="Elasmobranchii" family="Narcinidae" genus="Diplobatis" kingdom="Animalia" order="Torpediniformes" pageId="14" pageNumber="1202" phylum="Chordata" rank="genus">
<emphasis id="B935EA92227BB0512550EC15FE4AFABC" box="[290,403,1369,1392]" italics="true" pageId="14" pageNumber="1202">Diplobatis</emphasis>
</taxonomicName>
and
<taxonomicName id="4C414D03227BB05125A3EC15FDFEFABC" baseAuthorityName="Claeson" baseAuthorityYear="2014" box="[465,551,1369,1392]" class="Elasmobranchii" family="Narcinidae" genus="Narcine" kingdom="Animalia" order="Torpediniformes" pageId="14" pageNumber="1202" phylum="Chordata" rank="genus">
<emphasis id="B935EA92227BB05125A3EC15FDFEFABC" box="[465,551,1369,1392]" italics="true" pageId="14" pageNumber="1202">Narcine</emphasis>
</taxonomicName>
and is considered a derived trait of the narcinids among torpediniforms (see the phylogenetic analysis in this study).
</paragraph>
<caption id="DF3E6608227BB05124EDED59FBECFB59" ID-DOI="http://doi.org/10.5281/zenodo.10883311" ID-Zenodo-Dep="10883311" httpUri="https://zenodo.org/record/10883311/files/figure.png" pageId="14" pageNumber="1202" startId="14.[159,227,1045,1066]" targetBox="[173,1453,183,1013]" targetPageId="14" targetType="figure">
<paragraph id="8BFE3680227BB05124EDED59FBECFB59" blockId="14.[159,1467,1045,1173]" pageId="14" pageNumber="1202">
<emphasis id="B935EA92227BB05124EDED59FEB1FBE6" bold="true" box="[159,360,1045,1066]" pageId="14" pageNumber="1202">Figure 16. A, B, D,</emphasis>
<taxonomicName id="4C414D03227BB0512509ED59FD1FFBE6" authority="(Jaekel, 1894)" baseAuthorityName="Jaekel" baseAuthorityYear="1894" box="[379,710,1045,1066]" class="Elasmobranchii" family="Narcinidae" genus="Titanonarke" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="14" pageNumber="1202" phylum="Chordata" rank="species" species="molini">
<emphasis id="B935EA92227BB0512509ED59FDECFBE6" box="[379,565,1045,1066]" italics="true" pageId="14" pageNumber="1202">Titanonarke molini</emphasis>
(
<bibRefCitation id="EFD04B71227BB0512631ED59FD67FBE6" author="Jaekel, O." box="[579,702,1045,1066]" pageId="14" pageNumber="1202" refId="ref23357" refString="Jaekel, O. 1894. Die eoca ¨ nen Selachier vom Monte Bolca: ein Beitrag zur Morphogenie der Wirbelthiere. Springer, Berlin, 177 pp." type="book" year="1894">Jaekel, 1894</bibRefCitation>
)
</taxonomicName>
from the Eocene Monte Postale site;
<emphasis id="B935EA92227BB0512049ED59FB8AFBE6" bold="true" box="[1083,1107,1045,1066]" pageId="14" pageNumber="1202">A,</emphasis>
close-up of the abdominal region of MGP-PD 26275 showing the embryo; the anterior region of the body lies on the lower portion of the photo;
<emphasis id="B935EA92227BB05120C7ED7CFB13FB89" bold="true" box="[1205,1226,1072,1093]" pageId="14" pageNumber="1202">B,</emphasis>
reconstruction;
<emphasis id="B935EA92227BB0512115ED7CFAA6FB89" bold="true" box="[1383,1407,1072,1093]" pageId="14" pageNumber="1202">D,</emphasis>
detail of the vertebral column of the embryo indicated in B with a dotted rectangle.
<emphasis id="B935EA92227BB05127EDED07FC6FFBAC" bold="true" box="[927,950,1099,1120]" pageId="14" pageNumber="1202">C,</emphasis>
dissected specimen of
<taxonomicName id="4C414D03227BB05120EEED07FAB7FBAC" authorityName="Carvalho, Sabaj Perez &amp; Lovejoy" authorityYear="2011" box="[1180,1390,1099,1120]" class="Elasmobranchii" family="Potamotrygonidae" genus="Potamotrygon" kingdom="Animalia" order="Myliobatiformes" pageId="14" pageNumber="1202" phylum="Chordata" rank="species" species="tigrina">
<emphasis id="B935EA92227BB05120EEED07FAB7FBAC" box="[1180,1390,1099,1120]" italics="true" pageId="14" pageNumber="1202">Potamotrygon tigrina</emphasis>
</taxonomicName>
(IUWP 7361) showing the position of the left uterus, just next to the stomach. Abbreviations: int, intestine; liv, liver; lu, left uterus; na, neural arches; ru, right uterus; st, stomach; vc, vertebral centra. Scale bars: A, B = 10 mm; D = 1 mm.
</paragraph>
</caption>
<paragraph id="8BFE3680227BB05124B5EC94FB0AFA1C" blockId="14.[159,781,1496,1934]" lastBlockId="14.[845,1467,1241,1934]" pageId="14" pageNumber="1202">
<emphasis id="B935EA92227BB05124B5EC94FE0EFA23" bold="true" box="[199,471,1496,1519]" pageId="14" pageNumber="1202">Paired fins and girdles.</emphasis>
The scapulocoracoid is the largest element of the pectoral girdle. It is robust and strongly arched, and its anteriormost margin is situated between the basibranchial copula and the posterior tips of the synarcual flanges. The scapulocoracoid articulates anteriorly with the fifth pair of ceratobranchials. The suprascapulae are slender and fused medially, forming a slightly bowed bar in
<collectionCode id="ED50AE45227BB05125E1EFF4FE28F903" box="[403,497,1720,1743]" pageId="14" pageNumber="1202">MCSNV</collectionCode>
<collectionCode id="ED50AE45227BB0512588EFF4FDCEF903" box="[506,535,1720,1743]" country="China" name="Institute of Geology" pageId="14" pageNumber="1202">IG</collectionCode>
.VR.91359 (
<figureCitation id="137A2A05227BB05126EFEFF4FD07F903" box="[669,734,1720,1743]" captionStart="Figure 6" captionStartId="8.[159,227,1806,1827]" captionTargetBox="[365,1252,1347,1774]" captionTargetId="figure-317@8.[365,1261,1347,1774]" captionTargetPageId="8" captionText="Figure 6. †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site. A, the juvenile individual MCSNV IG.VR.91359; B, reconstruction. Abbreviations: ao, antorbital cartilage; cb, ceratobranchials; hym, hyomandibula; la, labial cartilages; me, Meckels cartilage; mes, mesopterygium; met, metapterygium; nc, nasal capsule; pel, prepelvic process; pub, puboischiadic bar; pq, palatoquadrate; pro, propterygium; ps, pseudohyoid; rad, pectoral radials; ro, rostral cartilage; sca, scapulocoracoid; ss, suprascapula; syn, synarcual. Scale bars = 50 mm." figureDoi="http://doi.org/10.5281/zenodo.10883285" httpUri="https://zenodo.org/record/10883285/files/figure.png" pageId="14" pageNumber="1202">Fig. 6</figureCitation>
). In
<collectionCode id="ED50AE45227BB05124EDEF94FF24F923" box="[159,253,1752,1775]" pageId="14" pageNumber="1202">MCSNV</collectionCode>
<collectionCode id="ED50AE45227BB051257FEF94FEF3F923" box="[269,298,1752,1775]" country="China" name="Institute of Geology" pageId="14" pageNumber="1202">IG</collectionCode>
.VR67290 the suprascapula appears to be inclined with respect to the vertebral column, confirming the interpretation of a bowed shape (
<figureCitation id="137A2A05227BB051264EEE5BFDA4F8E2" box="[572,637,1815,1838]" captionStart="Figure 5" captionStartId="8.[159,227,682,703]" captionTargetBox="[365,1255,183,650]" captionTargetId="figure-236@8.[365,1261,183,650]" captionTargetPageId="8" captionText="Figure 5. †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site. A, MCSNV IG.VR.67290, close-up of the head and hyoid apparatus; B, reconstruction. Abbreviations: ao, antorbital cartilage; bb, basibranchials; bbc, basibranchial copula; cb, ceratobranchials; cc, chondrocranium; eb, epibranchials; hb, hypobranchials; hym, hyomandibula; la, labial cartilages; me, Meckels cartilage; nc, nasal capsule; pq, palatoquadrate; ps, pseudohyoid; ra, rostral appendix; rf, rostral fontanelle; ro, rostral cartilage; sca, scapulocoracoid; ss, suprascapula; syn, synarcual. Scale bars = 50 mm." figureDoi="http://doi.org/10.5281/zenodo.10883279" httpUri="https://zenodo.org/record/10883279/files/figure.png" pageId="14" pageNumber="1202">Fig. 5</figureCitation>
). The suprascapular antimere is longer than the scapular process of the scapulocoracoid, which is posteriorly directed as in
<taxonomicName id="4C414D03227BB05124EDEE3BFF2CF842" baseAuthorityName="Claeson" baseAuthorityYear="2014" box="[159,245,1911,1934]" class="Elasmobranchii" family="Narcinidae" genus="Narcine" kingdom="Animalia" order="Torpediniformes" pageId="14" pageNumber="1202" phylum="Chordata" rank="genus">
<emphasis id="B935EA92227BB05124EDEE3BFF2CF842" box="[159,245,1911,1934]" italics="true" pageId="14" pageNumber="1202">Narcine</emphasis>
</taxonomicName>
(
<figureCitation id="137A2A05227BB0512577EE3BFE9BF842" box="[261,322,1911,1934]" captionStart="Figure 9" captionStartId="10.[159,227,702,723]" captionTargetBox="[365,1261,183,670]" captionTargetId="figure-180@10.[365,1261,183,670]" captionTargetPageId="10" captionText="Figure 9. A, synarcual and pectoral girdle of Narcine brasiliensis (AMNH 95343) in dorsal view; B, synarcual and pectoral girdle of †Titanonarke molini (MCSNV IG.VR.91359) in ventral view. The arrowheads indicate the posteriorly directed lateral stays of the synarcual. The arrows indicate the posteriorly directed scapular process of the scapulocoracoid. Abbreviations: cb, ceratocranchials; sca, scapulocoracoid; ss, suprascapula; syn, synarcual. Scale bar = 5 mm." figureDoi="http://doi.org/10.5281/zenodo.10883293" httpUri="https://zenodo.org/record/10883293/files/figure.png" pageId="14" pageNumber="1202">Fig. 9</figureCitation>
). The fusion of the suprascapular antimere with a visible suture is considered a synapomorphy of
<taxonomicName id="4C414D03227BB051273FEDB5FBD9FADC" authorityName="de Buen" authorityYear="1926" box="[845,1024,1273,1296]" class="Elasmobranchii" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="14" pageNumber="1202" phylum="Chordata" rank="order">Torpediniformes</taxonomicName>
by
<bibRefCitation id="EFD04B71227BB0512040EDB5FB05FADC" author="Claeson, K. M." box="[1074,1244,1273,1296]" pageId="14" pageNumber="1202" pagination="597 - 612" refId="ref20593" refString="Claeson, K. M. 2014. The impacts of comparative anatomy of electric rays (Batoidea: Torpediniformes) on their systematic hypotheses. Journal of Morphology, 275, 597 - 612." type="journal article" year="2014">Claeson (2014)</bibRefCitation>
. Although
<collectionCode id="ED50AE45227BB051212EEDB5FA63FADC" box="[1372,1466,1273,1296]" pageId="14" pageNumber="1202">MCSNV</collectionCode>
<collectionCode id="ED50AE45227BB051273FEC55FCB3FAFC" box="[845,874,1305,1328]" country="China" name="Institute of Geology" pageId="14" pageNumber="1202">IG</collectionCode>
.VR.67290 and
<collectionCode id="ED50AE45227BB0512052EC55FBA7FAFC" box="[1056,1150,1305,1328]" pageId="14" pageNumber="1202">MCSNV</collectionCode>
<collectionCode id="ED50AE45227BB05120FFEC55FB73FAFC" box="[1165,1194,1305,1328]" country="China" name="Institute of Geology" pageId="14" pageNumber="1202">IG</collectionCode>
.VR.91359 preserve this skeletal element (
<figureCitation id="137A2A05227BB051207BEC75FB93FA9C" box="[1033,1098,1337,1360]" captionStart="Figure 5" captionStartId="8.[159,227,682,703]" captionTargetBox="[365,1255,183,650]" captionTargetId="figure-236@8.[365,1261,183,650]" captionTargetPageId="8" captionText="Figure 5. †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site. A, MCSNV IG.VR.67290, close-up of the head and hyoid apparatus; B, reconstruction. Abbreviations: ao, antorbital cartilage; bb, basibranchials; bbc, basibranchial copula; cb, ceratobranchials; cc, chondrocranium; eb, epibranchials; hb, hypobranchials; hym, hyomandibula; la, labial cartilages; me, Meckels cartilage; nc, nasal capsule; pq, palatoquadrate; ps, pseudohyoid; ra, rostral appendix; rf, rostral fontanelle; ro, rostral cartilage; sca, scapulocoracoid; ss, suprascapula; syn, synarcual. Scale bars = 50 mm." figureDoi="http://doi.org/10.5281/zenodo.10883279" httpUri="https://zenodo.org/record/10883279/files/figure.png" pageId="14" pageNumber="1202">Figs 5</figureCitation>
,
<figureCitation id="137A2A05227BB0512025EC75FBBDFA9C" box="[1111,1124,1337,1360]" captionStart="Figure 6" captionStartId="8.[159,227,1806,1827]" captionTargetBox="[365,1252,1347,1774]" captionTargetId="figure-317@8.[365,1261,1347,1774]" captionTargetPageId="8" captionText="Figure 6. †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site. A, the juvenile individual MCSNV IG.VR.91359; B, reconstruction. Abbreviations: ao, antorbital cartilage; cb, ceratobranchials; hym, hyomandibula; la, labial cartilages; me, Meckels cartilage; mes, mesopterygium; met, metapterygium; nc, nasal capsule; pel, prepelvic process; pub, puboischiadic bar; pq, palatoquadrate; pro, propterygium; ps, pseudohyoid; rad, pectoral radials; ro, rostral cartilage; sca, scapulocoracoid; ss, suprascapula; syn, synarcual. Scale bars = 50 mm." figureDoi="http://doi.org/10.5281/zenodo.10883285" httpUri="https://zenodo.org/record/10883285/files/figure.png" pageId="14" pageNumber="1202">6</figureCitation>
,
<figureCitation id="137A2A05227BB0512003EC75FBA7FA9C" box="[1137,1150,1337,1360]" captionStart="Figure 9" captionStartId="10.[159,227,702,723]" captionTargetBox="[365,1261,183,670]" captionTargetId="figure-180@10.[365,1261,183,670]" captionTargetPageId="10" captionText="Figure 9. A, synarcual and pectoral girdle of Narcine brasiliensis (AMNH 95343) in dorsal view; B, synarcual and pectoral girdle of †Titanonarke molini (MCSNV IG.VR.91359) in ventral view. The arrowheads indicate the posteriorly directed lateral stays of the synarcual. The arrows indicate the posteriorly directed scapular process of the scapulocoracoid. Abbreviations: cb, ceratocranchials; sca, scapulocoracoid; ss, suprascapula; syn, synarcual. Scale bar = 5 mm." figureDoi="http://doi.org/10.5281/zenodo.10883293" httpUri="https://zenodo.org/record/10883293/files/figure.png" pageId="14" pageNumber="1202">9</figureCitation>
), it is not possible to describe a visible suture because both specimens are exposed ventrally and this area is obscured by the vertebral column. However, it is expected that †
<taxonomicName id="4C414D03227BB05120C0ECD5FAEDFA7C" authorityName="Carvalho" authorityYear="2010" box="[1202,1332,1433,1456]" class="Elasmobranchii" family="Narcinidae" genus="Titanonarke" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="14" pageNumber="1202" phylum="Chordata" rank="genus">
<emphasis id="B935EA92227BB05120C0ECD5FAEDFA7C" box="[1202,1332,1433,1456]" italics="true" pageId="14" pageNumber="1202">Titanonarke</emphasis>
</taxonomicName>
shares this character with all other electric rays.
</paragraph>
<paragraph id="8BFE3680227BB0512715EC94FB59F8E2" blockId="14.[845,1467,1241,1934]" pageId="14" pageNumber="1202">
The juvenile
<collectionCode id="ED50AE45227BB0512074EC94FBBDFA23" box="[1030,1124,1496,1519]" pageId="14" pageNumber="1202">MCSNV</collectionCode>
<collectionCode id="ED50AE45227BB0512006EC94FB4BFA23" box="[1140,1170,1496,1519]" country="China" name="Institute of Geology" pageId="14" pageNumber="1202">IG</collectionCode>
.VR.91359 shows a weak taphonomic displacement of the suprascapulae with respect to the vertebral column (
<figureCitation id="137A2A05227BB05120D5EF54FB31F9E3" box="[1191,1256,1560,1583]" captionStart="Figure 6" captionStartId="8.[159,227,1806,1827]" captionTargetBox="[365,1252,1347,1774]" captionTargetId="figure-317@8.[365,1261,1347,1774]" captionTargetPageId="8" captionText="Figure 6. †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site. A, the juvenile individual MCSNV IG.VR.91359; B, reconstruction. Abbreviations: ao, antorbital cartilage; cb, ceratobranchials; hym, hyomandibula; la, labial cartilages; me, Meckels cartilage; mes, mesopterygium; met, metapterygium; nc, nasal capsule; pel, prepelvic process; pub, puboischiadic bar; pq, palatoquadrate; pro, propterygium; ps, pseudohyoid; rad, pectoral radials; ro, rostral cartilage; sca, scapulocoracoid; ss, suprascapula; syn, synarcual. Scale bars = 50 mm." figureDoi="http://doi.org/10.5281/zenodo.10883285" httpUri="https://zenodo.org/record/10883285/files/figure.png" pageId="14" pageNumber="1202">Figs 6</figureCitation>
,
<figureCitation id="137A2A05227BB0512087EF54FADBF9E3" box="[1269,1282,1560,1583]" captionStart="Figure 9" captionStartId="10.[159,227,702,723]" captionTargetBox="[365,1261,183,670]" captionTargetId="figure-180@10.[365,1261,183,670]" captionTargetPageId="10" captionText="Figure 9. A, synarcual and pectoral girdle of Narcine brasiliensis (AMNH 95343) in dorsal view; B, synarcual and pectoral girdle of †Titanonarke molini (MCSNV IG.VR.91359) in ventral view. The arrowheads indicate the posteriorly directed lateral stays of the synarcual. The arrows indicate the posteriorly directed scapular process of the scapulocoracoid. Abbreviations: cb, ceratocranchials; sca, scapulocoracoid; ss, suprascapula; syn, synarcual. Scale bar = 5 mm." figureDoi="http://doi.org/10.5281/zenodo.10883293" httpUri="https://zenodo.org/record/10883293/files/figure.png" pageId="14" pageNumber="1202">9</figureCitation>
), which supports the hypothesis that the suprascapulae in torpediniforms are completely separate from the vertebral column and that the only connection between pectoral girdle and postcranium is through the fifth ceratobranchial (
<bibRefCitation id="EFD04B71227BB0512137EFD4FC08F903" author="Aschliman, N. C. &amp; Claeson, K. M. &amp; McEachran, J. D." pageId="14" pageNumber="1202" refId="ref18160" refString="Aschliman, N. C., Claeson, K. M. &amp; McEachran, J. D. 2012 a. Phylogeny of Batoidea. Pp. 57 - 96 in J. C. Carrier, J. A. Musick &amp; M. R. Heithaus (eds) Biology of sharks and their relatives. 2 nd edition. CRC Press, Boca Raton." type="book" year="2012">
Aschliman
<emphasis id="B935EA92227BB051273FEFF4FC59F903" box="[845,896,1720,1743]" italics="true" pageId="14" pageNumber="1202">et al</emphasis>
. 2012a
</bibRefCitation>
;
<bibRefCitation id="EFD04B71227BB0512796EFF4FBA0F903" author="Claeson, K. M." box="[996,1145,1720,1743]" pageId="14" pageNumber="1202" pagination="597 - 612" refId="ref20593" refString="Claeson, K. M. 2014. The impacts of comparative anatomy of electric rays (Batoidea: Torpediniformes) on their systematic hypotheses. Journal of Morphology, 275, 597 - 612." type="journal article" year="2014">Claeson 2014</bibRefCitation>
). The same specimen shows that the suprascapular projection of †
<taxonomicName id="4C414D03227BB05120AEEF94FA87F923" authorityName="Carvalho" authorityYear="2010" box="[1244,1374,1752,1775]" class="Elasmobranchii" family="Narcinidae" genus="Titanonarke" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="14" pageNumber="1202" phylum="Chordata" rank="genus">
<emphasis id="B935EA92227BB05120AEEF94FA87F923" box="[1244,1374,1752,1775]" italics="true" pageId="14" pageNumber="1202">Titanonarke</emphasis>
</taxonomicName>
was lateral and that the suprascapula-scapulocoracoid articulation was loose and unforked.
</paragraph>
<paragraph id="8BFE3680227BB04F2715EE7BFE56FBBD" blockId="14.[845,1467,1241,1934]" lastBlockId="16.[159,781,827,1934]" lastPageId="16" lastPageNumber="1204" pageId="14" pageNumber="1202">
The propterygia are long and arched and extend well beyond the anterior margin of the scapulocoracoid (
<figureCitation id="137A2A05227BB0512724EE3BFC7DF842" box="[854,932,1911,1934]" captionStart="Figure 10" captionStartId="10.[159,227,1833,1854]" captionTargetBox="[365,1248,1190,1801]" captionTargetId="figure-248@10.[365,1261,1190,1801]" captionTargetPageId="10" captionText="Figure 10. Pectoral fin of †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site. A, MCSNV IG.VR.67290; B, reconstruction; different colours are used to distinguish the propterygial (green), mesopterygial (yellow) and metapterygial (red) radials (colours in the online version). Abbreviations: mes, mesopterygium; met, metapterygium; pro, propterygium; sca, scapulocoracoid. Scale bars = 50 mm." figureDoi="http://doi.org/10.5281/zenodo.10883295" httpUri="https://zenodo.org/record/10883295/files/figure.png" pageId="14" pageNumber="1202">Fig. 10</figureCitation>
). They are composed of five or six propterygial segments. Two large voids, occupied in life by electric organs, are delimited by the propterygia, antorbital cartilages, hyoid archs and scapulocoracoids. The mesopterygium is flat, subtriangular in shape, and parallel and adjacent to the propterygium. The metapterygium is long and slender, is triangular in shape, and tapers posteriorly. The mesopterygium is shorter than the pro- and metapterygium in all specimens, although the apparent smaller size of the metapterygium in
<collectionCode id="ED50AE45227AB050267FEE3BFDB2F842" box="[525,619,1911,1934]" pageId="15" pageNumber="1203">MCSNV</collectionCode>
<collectionCode id="ED50AE45227AB050260EEE3BFD40F842" box="[636,665,1911,1934]" country="China" name="Institute of Geology" pageId="15" pageNumber="1203">IG</collectionCode>
.VR.67290 appears to be preservational (the metapterygium is weakly calcified in some narcinids; Carvalho &amp; Śeret 2002) or an artefact of the glue used to join pieces of the slab during preparation. There are about 40 pectoral radials (1216 propterygial, 912 mesopterygial and 1012 metapterygial), which bifurcate twice before reaching the edge of the pectoral fin margin. Each radial is composed of four segments before the first bifurcation, and another four elements before the second bifurcation (at least 910 segments in total;
<figureCitation id="137A2A052265B04F2510EA77FE74FC9E" box="[354,429,827,850]" captionStart="Figure 10" captionStartId="10.[159,227,1833,1854]" captionTargetBox="[365,1248,1190,1801]" captionTargetId="figure-248@10.[365,1261,1190,1801]" captionTargetPageId="10" captionText="Figure 10. Pectoral fin of †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site. A, MCSNV IG.VR.67290; B, reconstruction; different colours are used to distinguish the propterygial (green), mesopterygial (yellow) and metapterygial (red) radials (colours in the online version). Abbreviations: mes, mesopterygium; met, metapterygium; pro, propterygium; sca, scapulocoracoid. Scale bars = 50 mm." figureDoi="http://doi.org/10.5281/zenodo.10883295" httpUri="https://zenodo.org/record/10883295/files/figure.png" pageId="16" pageNumber="1204">Fig. 10</figureCitation>
). The lower number of segments recognized by
<bibRefCitation id="EFD04B712265B04F2531EA17FE2EFCBE" author="Carvalho, M. R. de" box="[323,503,859,882]" pageId="16" pageNumber="1204" refId="ref19921" refString="Carvalho, M. R. de. 2010. Morphology and phylogenetic relationships of the giant electric ray from the Eocene of Monte Bolca, Italy (Chondrichthyes: Torpediniformes). Pp. 183 - 198 in D. K. Elliot, J. G. Maisey, X. Yu &amp; D. Miao (eds) Morphology, phylogeny and paleobiogeography of fossil fishes. Verlag Dr. Friedrich Pfeil, Munich." type="book" year="2010">Carvalho (2010)</bibRefCitation>
, and actually detected in some specimens, is probably related to the loss of distal elements due to taphonomic processes. The radials are covered with a continuous layer of small (less than
<quantity id="4CB99B652265B04F26B4EAF7FCDFFC1E" box="[710,774,955,978]" metricMagnitude="-3" metricUnit="m" metricValue="1.0" pageId="16" pageNumber="1204" unit="mm" value="1.0">1 mm</quantity>
) tesserae, forming the so-called crustal calcification that characterizes the radials of basalmost batoids having an axial-undulatory swimming style, including pristids and rhinobatids, other than torpedinids and narcinids (Schaefer &amp; Summers 2005).
</paragraph>
<caption id="DF3E6608227AB05024EDE9F6FDF0FE6F" pageId="15" pageNumber="1203" startId="15.[159,217,186,207]" targetBox="[159,1454,448,1564]" targetIsTable="true" targetPageId="15" targetType="table">
<paragraph id="8BFE3680227AB05024EDE9F6FDF0FE6F" blockId="15.[159,1467,185,419]" pageId="15" pageNumber="1203">
<emphasis id="B935EA92227AB05024EDE9F6FF2DFF03" bold="true" box="[159,244,186,207]" pageId="15" pageNumber="1203">Table 1.</emphasis>
Morphometric data for all examined specimens of †
<taxonomicName id="4C414D03227AB050268EE9F6FBD7FF03" authority="Carvalho, 2010" authorityName="Carvalho" authorityYear="2010" box="[764,1038,186,207]" class="Elasmobranchii" family="Narcinidae" genus="Titanonarke" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="15" pageNumber="1203" phylum="Chordata" rank="genus">
<emphasis id="B935EA92227AB050268EE9F6FCA8FF03" box="[764,881,186,207]" italics="true" pageId="15" pageNumber="1203">Titanonarke</emphasis>
<bibRefCitation id="EFD04B71227AB050270BE9F6FBD7FF03" author="Carvalho, M. R. de" box="[889,1038,186,207]" pageId="15" pageNumber="1203" refId="ref19921" refString="Carvalho, M. R. de. 2010. Morphology and phylogenetic relationships of the giant electric ray from the Eocene of Monte Bolca, Italy (Chondrichthyes: Torpediniformes). Pp. 183 - 198 in D. K. Elliot, J. G. Maisey, X. Yu &amp; D. Miao (eds) Morphology, phylogeny and paleobiogeography of fossil fishes. Verlag Dr. Friedrich Pfeil, Munich." type="book" year="2010">Carvalho, 2010</bibRefCitation>
</taxonomicName>
from the Eocene Monte Postale site, Bolca Lagerst¨atte. Abbreviations (see the morphological scheme in the Supplemental material): AOW, antorbital cartilage width; CFL, caudal fin length; CFD, caudal fin depth; CLO, clasper length; CPD, caudal peduncle depth; D1B, first dorsal fin base length; D2B, second dorsal fin base length; DCS, dorsal caudal space; DL, disc length; DW, disc width; HL, head length; IDS, interdorsal space; MOW, mouth width; P2A, pelvic fin anterior margin length; P2B, pelvic fin base length; P2S, pelvic fin span; PCS, space from pelvic fin insertion to the caudal fin origin; PD1, predorsal distance up to the first dorsal fin; PD2, predorsal distance up to the second dorsal fin; PDI, pelvic fin insertion to first dorsal fin origin; PDO, pelvic fin origin to first dorsal fin origin; PGW, snout tip to the level of the greatest disc width; PIW, body width at pectoral fin insertions; POR, preoral length; PP2, prepelvic length; SL, standard length; TAL, tail length; TBW, tail base at pelvic fin origin; TL, total length.
</paragraph>
</caption>
<paragraph id="8BFE3680227AB0502621E897FA7BF9D0" pageId="15" pageNumber="1203">
<table id="F941C420227A4FA024EDE88CFA77F9D0" box="[159,1454,448,1564]" gridcols="13" gridrows="32" pageId="15" pageNumber="1203">
<tr id="357134C2227A4FA024EDE88CFA77FE3D" box="[159,1454,448,497]" gridrow="0" pageId="15" pageNumber="1203" rowspan-0="1" rowspan-1="1" rowspan-10="1" rowspan-2="1" rowspan-3="1" rowspan-9="1">
<th id="76A05DBE227A4FA02621E88CFC30FE3D" box="[595,1001,448,497]" colspan="5" colspanRight="4" gridcol="4" gridrow="0" pageId="15" pageNumber="1203">
<taxonomicName id="4C414D03227AB050262FE897FC7AFE3C" authority="(Jaekel, 1894)" baseAuthorityName="Jaekel" baseAuthorityYear="1894" box="[605,931,475,496]" class="Elasmobranchii" family="Narcinidae" genus="Titanonarke" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="15" pageNumber="1203" phylum="Chordata" rank="species" species="molini">
<emphasis id="B935EA92227AB050262FE897FCCCFE3C" box="[605,789,475,496]" italics="true" pageId="15" pageNumber="1203">Titanonarke molini</emphasis>
(
<bibRefCitation id="EFD04B71227AB0502750E897FC42FE3C" author="Jaekel, O." box="[802,923,475,496]" pageId="15" pageNumber="1203" refId="ref23357" refString="Jaekel, O. 1894. Die eoca ¨ nen Selachier vom Monte Bolca: ein Beitrag zur Morphogenie der Wirbelthiere. Springer, Berlin, 177 pp." type="book" year="1894">Jaekel, 1894</bibRefCitation>
)
</taxonomicName>
</th>
<th id="76A05DBE227A4FA020AAE88CFA77FE3D" box="[1240,1454,448,497]" colspan="2" colspanRight="1" gridcol="11" gridrow="0" pageId="15" pageNumber="1203">
<taxonomicName id="4C414D03227AB050217FE88DFA85FE3C" authority="Marram̀ &amp; Claeso &amp; Carneval &amp; Kriwe, 2018" authorityName="Marram̀ &amp; Claeso &amp; Carneval &amp; Kriwe" authorityYear="2018" class="Elasmobranchii" family="Narcinidae" genus="Titanonarke" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="15" pageNumber="1203" phylum="Chordata" rank="species" species="megapterygia" status="sp. nov.">
<emphasis id="B935EA92227AB050217FE88DFA85FE3C" italics="true" pageId="15" pageNumber="1203">Titanonarke megapterygia</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A20657E9227AB0502110E897FA77FE3C" box="[1378,1454,475,496]" pageId="15" pageNumber="1203" rank="species">sp. nov.</taxonomicNameLabel>
</th>
</tr>
<tr id="357134C2227A4FA024EDEB41FA77FDF0" box="[159,1454,525,572]" gridrow="1" pageId="15" pageNumber="1203" rowspan-0="1">
<td id="76A05DBE227A4FA0253EEB41FE00FDF0" box="[332,473,525,572]" colspan="2" colspanRight="1" gridcol="1" gridrow="1" pageId="15" pageNumber="1203">MGP-PD 26275/6</td>
<td id="76A05DBE227A4FA0258DEB41FD4DFDF0" box="[511,660,525,572]" colspan="2" colspanRight="1" gridcol="3" gridrow="1" pageId="15" pageNumber="1203">MCSNV IG.VR.67290</td>
<td id="76A05DBE227A4FA026CBEB41FC9FFDF0" box="[697,838,525,572]" colspan="2" colspanRight="1" gridcol="5" gridrow="1" pageId="15" pageNumber="1203">MCSNV IG.91128/9</td>
<td id="76A05DBE227A4FA02715EB41FC30FDF0" box="[871,1001,525,572]" colspan="2" colspanRight="1" gridcol="7" gridrow="1" pageId="15" pageNumber="1203">MCSNV IG.135581</td>
<td id="76A05DBE227A4FA02061EB41FB7BFDF0" box="[1043,1186,525,572]" colspan="2" colspanRight="1" gridcol="9" gridrow="1" pageId="15" pageNumber="1203">MCSNV IG.VR.91359</td>
<td id="76A05DBE227A4FA020AAEB41FA77FDF0" box="[1240,1454,525,572]" colspan="2" colspanRight="1" gridcol="11" gridrow="1" pageId="15" pageNumber="1203">MCSNV IG.135576</td>
</tr>
<tr id="357134C2227A4FA024EDEB15FA77FDA2" box="[159,1454,601,622]" gridrow="2" pageId="15" pageNumber="1203">
<th id="76A05DBE227A4FA024EDEB15FEF2FDA2" box="[159,299,601,622]" gridcol="0" gridrow="2" pageId="15" pageNumber="1203">Measurements</th>
<td id="76A05DBE227A4FA0253EEB15FE5BFDA2" box="[332,386,601,622]" gridcol="1" gridrow="2" pageId="15" pageNumber="1203">mm</td>
<td id="76A05DBE227A4FA025D1EB15FE00FDA2" box="[419,473,601,622]" gridcol="2" gridrow="2" pageId="15" pageNumber="1203">% SL</td>
<td id="76A05DBE227A4FA0258DEB15FDECFDA2" box="[511,565,601,622]" gridcol="3" gridrow="2" pageId="15" pageNumber="1203">mm</td>
<td id="76A05DBE227A4FA02621EB15FD4DFDA2" box="[595,660,601,622]" gridcol="4" gridrow="2" pageId="15" pageNumber="1203">% SL</td>
<td id="76A05DBE227A4FA026CBEB15FD36FDA2" box="[697,751,601,622]" gridcol="5" gridrow="2" pageId="15" pageNumber="1203">mm</td>
<td id="76A05DBE227A4FA02762EB15FC9FFDA2" box="[784,838,601,622]" gridcol="6" gridrow="2" pageId="15" pageNumber="1203">% SL</td>
<td id="76A05DBE227A4FA02715EB15FC30FDA2" box="[871,1001,601,622]" colspan="2" colspanRight="1" gridcol="7" gridrow="2" pageId="15" pageNumber="1203">mm % SL</td>
<td id="76A05DBE227A4FA02061EB15FBE4FDA2" box="[1043,1085,601,622]" gridcol="9" gridrow="2" pageId="15" pageNumber="1203">mm</td>
<td id="76A05DBE227A4FA02017EB15FB7BFDA2" box="[1125,1186,601,622]" gridcol="10" gridrow="2" pageId="15" pageNumber="1203">% SL</td>
<td id="76A05DBE227A4FA020AAEB15FAC0FDA2" box="[1240,1305,601,622]" gridcol="11" gridrow="2" pageId="15" pageNumber="1203">mm</td>
<td id="76A05DBE227A4FA0211EEB15FA77FDA2" box="[1388,1454,601,622]" gridcol="12" gridrow="2" pageId="15" pageNumber="1203">% SL</td>
</tr>
<tr id="357134C2227A4FA024EDEBC6FA77FD53" box="[159,1454,650,671]" gridrow="3" pageId="15" pageNumber="1203">
<th id="76A05DBE227A4FA024EDEBC6FEF2FD53" box="[159,299,650,671]" gridcol="0" gridrow="3" pageId="15" pageNumber="1203">AOW</th>
<td id="76A05DBE227A4FA0253EEBC6FE5BFD53" box="[332,386,650,671]" gridcol="1" gridrow="3" pageId="15" pageNumber="1203">206.6</td>
<td id="76A05DBE227A4FA025D1EBC6FE00FD53" box="[419,473,650,671]" gridcol="2" gridrow="3" pageId="15" pageNumber="1203">25.3</td>
<td id="76A05DBE227A4FA0258DEBC6FDECFD53" box="[511,565,650,671]" gridcol="3" gridrow="3" pageId="15" pageNumber="1203">217.4</td>
<td id="76A05DBE227A4FA02621EBC6FD4DFD53" box="[595,660,650,671]" gridcol="4" gridrow="3" pageId="15" pageNumber="1203">29.0</td>
<td id="76A05DBE227A4FA026CBEBC6FD36FD53" box="[697,751,650,671]" gridcol="5" gridrow="3" pageId="15" pageNumber="1203">196.2</td>
<td id="76A05DBE227A4FA02762EBC6FC9FFD53" box="[784,838,650,671]" gridcol="6" gridrow="3" pageId="15" pageNumber="1203">23.1</td>
<td id="76A05DBE227A4FA02715EBC6FC4BFD53" box="[871,914,650,671]" gridcol="7" gridrow="3" pageId="15" pageNumber="1203">26.1</td>
<td id="76A05DBE227A4FA027BBEBC6FC30FD53" box="[969,1001,650,671]" gridcol="8" gridrow="3" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02061EBC6FBE4FD53" box="[1043,1085,650,671]" gridcol="9" gridrow="3" pageId="15" pageNumber="1203">28.1</td>
<td id="76A05DBE227A4FA02017EBC6FB7BFD53" box="[1125,1186,650,671]" gridcol="10" gridrow="3" pageId="15" pageNumber="1203">28.4</td>
<td id="76A05DBE227A4FA020AAEBC6FAC0FD53" box="[1240,1305,650,671]" gridcol="11" gridrow="3" pageId="15" pageNumber="1203">174.2</td>
<td id="76A05DBE227A4FA0211EEBC6FA77FD53" box="[1388,1454,650,671]" gridcol="12" gridrow="3" pageId="15" pageNumber="1203">27.8</td>
</tr>
<tr id="357134C2227A4FA024EDEBE6FA77FD73" box="[159,1454,682,703]" gridrow="4" pageId="15" pageNumber="1203">
<th id="76A05DBE227A4FA024EDEBE6FEF2FD73" box="[159,299,682,703]" gridcol="0" gridrow="4" pageId="15" pageNumber="1203">CFL</th>
<td id="76A05DBE227A4FA0253EEBE6FE5BFD73" box="[332,386,682,703]" gridcol="1" gridrow="4" pageId="15" pageNumber="1203">80.9</td>
<td id="76A05DBE227A4FA025D1EBE6FE00FD73" box="[419,473,682,703]" gridcol="2" gridrow="4" pageId="15" pageNumber="1203">9.9</td>
<td id="76A05DBE227A4FA0258DEBE6FDECFD73" box="[511,565,682,703]" gridcol="3" gridrow="4" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02621EBE6FD4DFD73" box="[595,660,682,703]" gridcol="4" gridrow="4" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA026CBEBE6FD36FD73" box="[697,751,682,703]" gridcol="5" gridrow="4" pageId="15" pageNumber="1203">75.0</td>
<td id="76A05DBE227A4FA02762EBE6FC9FFD73" box="[784,838,682,703]" gridcol="6" gridrow="4" pageId="15" pageNumber="1203">8.8</td>
<td id="76A05DBE227A4FA02715EBE6FC4BFD73" box="[871,914,682,703]" gridcol="7" gridrow="4" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA027BBEBE6FC30FD73" box="[969,1001,682,703]" gridcol="8" gridrow="4" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02061EBE6FBE4FD73" box="[1043,1085,682,703]" gridcol="9" gridrow="4" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02017EBE6FB7BFD73" box="[1125,1186,682,703]" gridcol="10" gridrow="4" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA020AAEBE6FAC0FD73" box="[1240,1305,682,703]" gridcol="11" gridrow="4" pageId="15" pageNumber="1203">97.4</td>
<td id="76A05DBE227A4FA0211EEBE6FA77FD73" box="[1388,1454,682,703]" gridcol="12" gridrow="4" pageId="15" pageNumber="1203">15.6</td>
</tr>
<tr id="357134C2227A4FA024EDEB86FA77FD13" box="[159,1454,714,735]" gridrow="5" pageId="15" pageNumber="1203">
<th id="76A05DBE227A4FA024EDEB86FEF2FD13" box="[159,299,714,735]" gridcol="0" gridrow="5" pageId="15" pageNumber="1203">CFD</th>
<td id="76A05DBE227A4FA0253EEB86FE5BFD13" box="[332,386,714,735]" gridcol="1" gridrow="5" pageId="15" pageNumber="1203">28.4</td>
<td id="76A05DBE227A4FA025D1EB86FE00FD13" box="[419,473,714,735]" gridcol="2" gridrow="5" pageId="15" pageNumber="1203">3.5</td>
<td id="76A05DBE227A4FA0258DEB86FDECFD13" box="[511,565,714,735]" gridcol="3" gridrow="5" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02621EB86FD4DFD13" box="[595,660,714,735]" gridcol="4" gridrow="5" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA026CBEB86FD36FD13" box="[697,751,714,735]" gridcol="5" gridrow="5" pageId="15" pageNumber="1203">26.1</td>
<td id="76A05DBE227A4FA02762EB86FC9FFD13" box="[784,838,714,735]" gridcol="6" gridrow="5" pageId="15" pageNumber="1203">3.1</td>
<td id="76A05DBE227A4FA02715EB86FC4BFD13" box="[871,914,714,735]" gridcol="7" gridrow="5" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA027BBEB86FC30FD13" box="[969,1001,714,735]" gridcol="8" gridrow="5" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02061EB86FBE4FD13" box="[1043,1085,714,735]" gridcol="9" gridrow="5" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02017EB86FB7BFD13" box="[1125,1186,714,735]" gridcol="10" gridrow="5" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA020AAEB86FAC0FD13" box="[1240,1305,714,735]" gridcol="11" gridrow="5" pageId="15" pageNumber="1203">36.0</td>
<td id="76A05DBE227A4FA0211EEB86FA77FD13" box="[1388,1454,714,735]" gridcol="12" gridrow="5" pageId="15" pageNumber="1203">5.8</td>
</tr>
<tr id="357134C2227A4FA024EDEBA6FA77FD33" box="[159,1454,746,767]" gridrow="6" pageId="15" pageNumber="1203">
<th id="76A05DBE227A4FA024EDEBA6FEF2FD33" box="[159,299,746,767]" gridcol="0" gridrow="6" pageId="15" pageNumber="1203">CLO</th>
<td id="76A05DBE227A4FA0253EEBA6FE5BFD33" box="[332,386,746,767]" gridcol="1" gridrow="6" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA025D1EBA6FE00FD33" box="[419,473,746,767]" gridcol="2" gridrow="6" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA0258DEBA6FDECFD33" box="[511,565,746,767]" gridcol="3" gridrow="6" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02621EBA6FD4DFD33" box="[595,660,746,767]" gridcol="4" gridrow="6" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA026CBEBA6FD36FD33" box="[697,751,746,767]" gridcol="5" gridrow="6" pageId="15" pageNumber="1203">70.5</td>
<td id="76A05DBE227A4FA02762EBA6FC9FFD33" box="[784,838,746,767]" gridcol="6" gridrow="6" pageId="15" pageNumber="1203">8.3</td>
<td id="76A05DBE227A4FA02715EBA6FC4BFD33" box="[871,914,746,767]" gridcol="7" gridrow="6" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA027BBEBA6FC30FD33" box="[969,1001,746,767]" gridcol="8" gridrow="6" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02061EBA6FBE4FD33" box="[1043,1085,746,767]" gridcol="9" gridrow="6" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02017EBA6FB7BFD33" box="[1125,1186,746,767]" gridcol="10" gridrow="6" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA020AAEBA6FAC0FD33" box="[1240,1305,746,767]" gridcol="11" gridrow="6" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA0211EEBA6FA77FD33" box="[1388,1454,746,767]" gridcol="12" gridrow="6" pageId="15" pageNumber="1203"></td>
</tr>
<tr id="357134C2227A4FA024EDEA46FA77FCD3" box="[159,1454,778,799]" gridrow="7" pageId="15" pageNumber="1203">
<th id="76A05DBE227A4FA024EDEA46FEF2FCD3" box="[159,299,778,799]" gridcol="0" gridrow="7" pageId="15" pageNumber="1203">CPD</th>
<td id="76A05DBE227A4FA0253EEA46FE5BFCD3" box="[332,386,778,799]" gridcol="1" gridrow="7" pageId="15" pageNumber="1203">20.2</td>
<td id="76A05DBE227A4FA025D1EA46FE00FCD3" box="[419,473,778,799]" gridcol="2" gridrow="7" pageId="15" pageNumber="1203">2.5</td>
<td id="76A05DBE227A4FA0258DEA46FDECFCD3" box="[511,565,778,799]" gridcol="3" gridrow="7" pageId="15" pageNumber="1203">25.5</td>
<td id="76A05DBE227A4FA02621EA46FD4DFCD3" box="[595,660,778,799]" gridcol="4" gridrow="7" pageId="15" pageNumber="1203">3.4</td>
<td id="76A05DBE227A4FA026CBEA46FD36FCD3" box="[697,751,778,799]" gridcol="5" gridrow="7" pageId="15" pageNumber="1203">15.9</td>
<td id="76A05DBE227A4FA02762EA46FC9FFCD3" box="[784,838,778,799]" gridcol="6" gridrow="7" pageId="15" pageNumber="1203">1.9</td>
<td id="76A05DBE227A4FA02715EA46FC4BFCD3" box="[871,914,778,799]" gridcol="7" gridrow="7" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA027BBEA46FC30FCD3" box="[969,1001,778,799]" gridcol="8" gridrow="7" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02061EA46FBE4FCD3" box="[1043,1085,778,799]" gridcol="9" gridrow="7" pageId="15" pageNumber="1203">2.3</td>
<td id="76A05DBE227A4FA02017EA46FB7BFCD3" box="[1125,1186,778,799]" gridcol="10" gridrow="7" pageId="15" pageNumber="1203">2.3</td>
<td id="76A05DBE227A4FA020AAEA46FAC0FCD3" box="[1240,1305,778,799]" gridcol="11" gridrow="7" pageId="15" pageNumber="1203">13.7</td>
<td id="76A05DBE227A4FA0211EEA46FA77FCD3" box="[1388,1454,778,799]" gridcol="12" gridrow="7" pageId="15" pageNumber="1203">2.2</td>
</tr>
<tr id="357134C2227A4FA024EDEA66FA77FCF3" box="[159,1454,810,831]" gridrow="8" pageId="15" pageNumber="1203">
<th id="76A05DBE227A4FA024EDEA66FEF2FCF3" box="[159,299,810,831]" gridcol="0" gridrow="8" pageId="15" pageNumber="1203">D1B</th>
<td id="76A05DBE227A4FA0253EEA66FE5BFCF3" box="[332,386,810,831]" gridcol="1" gridrow="8" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA025D1EA66FE00FCF3" box="[419,473,810,831]" gridcol="2" gridrow="8" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA0258DEA66FDECFCF3" box="[511,565,810,831]" gridcol="3" gridrow="8" pageId="15" pageNumber="1203">64.6</td>
<td id="76A05DBE227A4FA02621EA66FD4DFCF3" box="[595,660,810,831]" gridcol="4" gridrow="8" pageId="15" pageNumber="1203">8.6</td>
<td id="76A05DBE227A4FA026CBEA66FD36FCF3" box="[697,751,810,831]" gridcol="5" gridrow="8" pageId="15" pageNumber="1203">56.6</td>
<td id="76A05DBE227A4FA02762EA66FC9FFCF3" box="[784,838,810,831]" gridcol="6" gridrow="8" pageId="15" pageNumber="1203">6.7</td>
<td id="76A05DBE227A4FA02715EA66FC4BFCF3" box="[871,914,810,831]" gridcol="7" gridrow="8" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA027BBEA66FC30FCF3" box="[969,1001,810,831]" gridcol="8" gridrow="8" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02061EA66FBE4FCF3" box="[1043,1085,810,831]" gridcol="9" gridrow="8" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02017EA66FB7BFCF3" box="[1125,1186,810,831]" gridcol="10" gridrow="8" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA020AAEA66FAC0FCF3" box="[1240,1305,810,831]" gridcol="11" gridrow="8" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA0211EEA66FA77FCF3" box="[1388,1454,810,831]" gridcol="12" gridrow="8" pageId="15" pageNumber="1203"></td>
</tr>
<tr id="357134C2227A4FA024EDEA06FA77FC93" box="[159,1454,842,863]" gridrow="9" pageId="15" pageNumber="1203">
<th id="76A05DBE227A4FA024EDEA06FEF2FC93" box="[159,299,842,863]" gridcol="0" gridrow="9" pageId="15" pageNumber="1203">D2B</th>
<td id="76A05DBE227A4FA0253EEA06FE5BFC93" box="[332,386,842,863]" gridcol="1" gridrow="9" pageId="15" pageNumber="1203">50.0</td>
<td id="76A05DBE227A4FA025D1EA06FE00FC93" box="[419,473,842,863]" gridcol="2" gridrow="9" pageId="15" pageNumber="1203">6.1</td>
<td id="76A05DBE227A4FA0258DEA06FDECFC93" box="[511,565,842,863]" gridcol="3" gridrow="9" pageId="15" pageNumber="1203">47.9</td>
<td id="76A05DBE227A4FA02621EA06FD4DFC93" box="[595,660,842,863]" gridcol="4" gridrow="9" pageId="15" pageNumber="1203">6.4</td>
<td id="76A05DBE227A4FA026CBEA06FD36FC93" box="[697,751,842,863]" gridcol="5" gridrow="9" pageId="15" pageNumber="1203">51.3</td>
<td id="76A05DBE227A4FA02762EA06FC9FFC93" box="[784,838,842,863]" gridcol="6" gridrow="9" pageId="15" pageNumber="1203">6.0</td>
<td id="76A05DBE227A4FA02715EA06FC4BFC93" box="[871,914,842,863]" gridcol="7" gridrow="9" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA027BBEA06FC30FC93" box="[969,1001,842,863]" gridcol="8" gridrow="9" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02061EA06FBE4FC93" box="[1043,1085,842,863]" gridcol="9" gridrow="9" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02017EA06FB7BFC93" box="[1125,1186,842,863]" gridcol="10" gridrow="9" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA020AAEA06FAC0FC93" box="[1240,1305,842,863]" gridcol="11" gridrow="9" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA0211EEA06FA77FC93" box="[1388,1454,842,863]" gridcol="12" gridrow="9" pageId="15" pageNumber="1203"></td>
</tr>
<tr id="357134C2227A4FA024EDEA25FA77FCB2" box="[159,1454,873,894]" gridrow="10" pageId="15" pageNumber="1203">
<th id="76A05DBE227A4FA024EDEA25FEF2FCB2" box="[159,299,873,894]" gridcol="0" gridrow="10" pageId="15" pageNumber="1203">DCS</th>
<td id="76A05DBE227A4FA0253EEA25FE5BFCB2" box="[332,386,873,894]" gridcol="1" gridrow="10" pageId="15" pageNumber="1203">140.2</td>
<td id="76A05DBE227A4FA025D1EA25FE00FCB2" box="[419,473,873,894]" gridcol="2" gridrow="10" pageId="15" pageNumber="1203">17.2</td>
<td id="76A05DBE227A4FA0258DEA25FDECFCB2" box="[511,565,873,894]" gridcol="3" gridrow="10" pageId="15" pageNumber="1203">109.8</td>
<td id="76A05DBE227A4FA02621EA25FD4DFCB2" box="[595,660,873,894]" gridcol="4" gridrow="10" pageId="15" pageNumber="1203">14.6</td>
<td id="76A05DBE227A4FA026CBEA25FD36FCB2" box="[697,751,873,894]" gridcol="5" gridrow="10" pageId="15" pageNumber="1203">119.3</td>
<td id="76A05DBE227A4FA02762EA25FC9FFCB2" box="[784,838,873,894]" gridcol="6" gridrow="10" pageId="15" pageNumber="1203">14.0</td>
<td id="76A05DBE227A4FA02715EA25FC4BFCB2" box="[871,914,873,894]" gridcol="7" gridrow="10" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA027BBEA25FC30FCB2" box="[969,1001,873,894]" gridcol="8" gridrow="10" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02061EA25FBE4FCB2" box="[1043,1085,873,894]" gridcol="9" gridrow="10" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02017EA25FB7BFCB2" box="[1125,1186,873,894]" gridcol="10" gridrow="10" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA020AAEA25FAC0FCB2" box="[1240,1305,873,894]" gridcol="11" gridrow="10" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA0211EEA25FA77FCB2" box="[1388,1454,873,894]" gridcol="12" gridrow="10" pageId="15" pageNumber="1203"></td>
</tr>
<tr id="357134C2227A4FA024EDEAC5FA77FC52" box="[159,1454,905,926]" gridrow="11" pageId="15" pageNumber="1203">
<th id="76A05DBE227A4FA024EDEAC5FEF2FC52" box="[159,299,905,926]" gridcol="0" gridrow="11" pageId="15" pageNumber="1203">DL</th>
<td id="76A05DBE227A4FA0253EEAC5FE5BFC52" box="[332,386,905,926]" gridcol="1" gridrow="11" pageId="15" pageNumber="1203">353.1</td>
<td id="76A05DBE227A4FA025D1EAC5FE00FC52" box="[419,473,905,926]" gridcol="2" gridrow="11" pageId="15" pageNumber="1203">43.2</td>
<td id="76A05DBE227A4FA0258DEAC5FDECFC52" box="[511,565,905,926]" gridcol="3" gridrow="11" pageId="15" pageNumber="1203">341.2</td>
<td id="76A05DBE227A4FA02621EAC5FD4DFC52" box="[595,660,905,926]" gridcol="4" gridrow="11" pageId="15" pageNumber="1203">45.5</td>
<td id="76A05DBE227A4FA026CBEAC5FD36FC52" box="[697,751,905,926]" gridcol="5" gridrow="11" pageId="15" pageNumber="1203">371.6</td>
<td id="76A05DBE227A4FA02762EAC5FC9FFC52" box="[784,838,905,926]" gridcol="6" gridrow="11" pageId="15" pageNumber="1203">43.7</td>
<td id="76A05DBE227A4FA02715EAC5FC4BFC52" box="[871,914,905,926]" gridcol="7" gridrow="11" pageId="15" pageNumber="1203">33.8</td>
<td id="76A05DBE227A4FA027BBEAC5FC30FC52" box="[969,1001,905,926]" gridcol="8" gridrow="11" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02061EAC5FBE4FC52" box="[1043,1085,905,926]" gridcol="9" gridrow="11" pageId="15" pageNumber="1203">42.8</td>
<td id="76A05DBE227A4FA02017EAC5FB7BFC52" box="[1125,1186,905,926]" gridcol="10" gridrow="11" pageId="15" pageNumber="1203">43.2</td>
<td id="76A05DBE227A4FA020AAEAC5FAC0FC52" box="[1240,1305,905,926]" gridcol="11" gridrow="11" pageId="15" pageNumber="1203">334.5</td>
<td id="76A05DBE227A4FA0211EEAC5FA77FC52" box="[1388,1454,905,926]" gridcol="12" gridrow="11" pageId="15" pageNumber="1203">53.4</td>
</tr>
<tr id="357134C2227A4FA024EDEAE5FA77FC72" box="[159,1454,937,958]" gridrow="12" pageId="15" pageNumber="1203">
<th id="76A05DBE227A4FA024EDEAE5FEF2FC72" box="[159,299,937,958]" gridcol="0" gridrow="12" pageId="15" pageNumber="1203">DW</th>
<td id="76A05DBE227A4FA0253EEAE5FE5BFC72" box="[332,386,937,958]" gridcol="1" gridrow="12" pageId="15" pageNumber="1203">387.0</td>
<td id="76A05DBE227A4FA025D1EAE5FE00FC72" box="[419,473,937,958]" gridcol="2" gridrow="12" pageId="15" pageNumber="1203">47.4</td>
<td id="76A05DBE227A4FA0258DEAE5FDECFC72" box="[511,565,937,958]" gridcol="3" gridrow="12" pageId="15" pageNumber="1203">406.6</td>
<td id="76A05DBE227A4FA02621EAE5FD4DFC72" box="[595,660,937,958]" gridcol="4" gridrow="12" pageId="15" pageNumber="1203">54.2</td>
<td id="76A05DBE227A4FA026CBEAE5FD36FC72" box="[697,751,937,958]" gridcol="5" gridrow="12" pageId="15" pageNumber="1203">347.8</td>
<td id="76A05DBE227A4FA02762EAE5FC9FFC72" box="[784,838,937,958]" gridcol="6" gridrow="12" pageId="15" pageNumber="1203">40.9</td>
<td id="76A05DBE227A4FA02715EAE5FC4BFC72" box="[871,914,937,958]" gridcol="7" gridrow="12" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA027BBEAE5FC30FC72" box="[969,1001,937,958]" gridcol="8" gridrow="12" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02061EAE5FBE4FC72" box="[1043,1085,937,958]" gridcol="9" gridrow="12" pageId="15" pageNumber="1203">56.3</td>
<td id="76A05DBE227A4FA02017EAE5FB7BFC72" box="[1125,1186,937,958]" gridcol="10" gridrow="12" pageId="15" pageNumber="1203">56.9</td>
<td id="76A05DBE227A4FA020AAEAE5FAC0FC72" box="[1240,1305,937,958]" gridcol="11" gridrow="12" pageId="15" pageNumber="1203">349.8</td>
<td id="76A05DBE227A4FA0211EEAE5FA77FC72" box="[1388,1454,937,958]" gridcol="12" gridrow="12" pageId="15" pageNumber="1203">55.9</td>
</tr>
<tr id="357134C2227A4FA024EDEA85FA77FC12" box="[159,1454,969,990]" gridrow="13" pageId="15" pageNumber="1203">
<th id="76A05DBE227A4FA024EDEA85FEF2FC12" box="[159,299,969,990]" gridcol="0" gridrow="13" pageId="15" pageNumber="1203">HL</th>
<td id="76A05DBE227A4FA0253EEA85FE5BFC12" box="[332,386,969,990]" gridcol="1" gridrow="13" pageId="15" pageNumber="1203">178.1</td>
<td id="76A05DBE227A4FA025D1EA85FE00FC12" box="[419,473,969,990]" gridcol="2" gridrow="13" pageId="15" pageNumber="1203">21.8</td>
<td id="76A05DBE227A4FA0258DEA85FDECFC12" box="[511,565,969,990]" gridcol="3" gridrow="13" pageId="15" pageNumber="1203">172.5</td>
<td id="76A05DBE227A4FA02621EA85FD4DFC12" box="[595,660,969,990]" gridcol="4" gridrow="13" pageId="15" pageNumber="1203">23.0</td>
<td id="76A05DBE227A4FA026CBEA85FD36FC12" box="[697,751,969,990]" gridcol="5" gridrow="13" pageId="15" pageNumber="1203">204.4</td>
<td id="76A05DBE227A4FA02762EA85FC9FFC12" box="[784,838,969,990]" gridcol="6" gridrow="13" pageId="15" pageNumber="1203">24.0</td>
<td id="76A05DBE227A4FA02715EA85FC4BFC12" box="[871,914,969,990]" gridcol="7" gridrow="13" pageId="15" pageNumber="1203">22.2</td>
<td id="76A05DBE227A4FA027BBEA85FC30FC12" box="[969,1001,969,990]" gridcol="8" gridrow="13" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02061EA85FBE4FC12" box="[1043,1085,969,990]" gridcol="9" gridrow="13" pageId="15" pageNumber="1203">23.6</td>
<td id="76A05DBE227A4FA02017EA85FB7BFC12" box="[1125,1186,969,990]" gridcol="10" gridrow="13" pageId="15" pageNumber="1203">23.8</td>
<td id="76A05DBE227A4FA020AAEA85FAC0FC12" box="[1240,1305,969,990]" gridcol="11" gridrow="13" pageId="15" pageNumber="1203">171.8</td>
<td id="76A05DBE227A4FA0211EEA85FA77FC12" box="[1388,1454,969,990]" gridcol="12" gridrow="13" pageId="15" pageNumber="1203">27.4</td>
</tr>
<tr id="357134C2227A4FA024EDEAA5FA77FC32" box="[159,1454,1001,1022]" gridrow="14" pageId="15" pageNumber="1203" rowspan-12="1">
<th id="76A05DBE227A4FA024EDEAA5FEF2FC32" box="[159,299,1001,1022]" gridcol="0" gridrow="14" pageId="15" pageNumber="1203">IDS</th>
<td id="76A05DBE227A4FA0253EEAA5FE5BFC32" box="[332,386,1001,1022]" gridcol="1" gridrow="14" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA025D1EAA5FE00FC32" box="[419,473,1001,1022]" gridcol="2" gridrow="14" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA0258DEAA5FDECFC32" box="[511,565,1001,1022]" gridcol="3" gridrow="14" pageId="15" pageNumber="1203">53.6</td>
<td id="76A05DBE227A4FA02621EAA5FD4DFC32" box="[595,660,1001,1022]" gridcol="4" gridrow="14" pageId="15" pageNumber="1203">7.1</td>
<td id="76A05DBE227A4FA026CBEAA5FD36FC32" box="[697,751,1001,1022]" gridcol="5" gridrow="14" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02762EAA5FC9FFC32" box="[784,838,1001,1022]" gridcol="6" gridrow="14" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02715EAA5FC4BFC32" box="[871,914,1001,1022]" gridcol="7" gridrow="14" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA027BBEAA5FC30FC32" box="[969,1001,1001,1022]" gridcol="8" gridrow="14" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02061EAA5FBE4FC32" box="[1043,1085,1001,1022]" gridcol="9" gridrow="14" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02017EAA5FB7BFC32" box="[1125,1186,1001,1022]" gridcol="10" gridrow="14" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA020AAEAA5FAC0FC32" box="[1240,1305,1001,1022]" gridcol="11" gridrow="14" pageId="15" pageNumber="1203"></td>
</tr>
<tr id="357134C2227A4FA024EDED45FA77FBD2" box="[159,1454,1033,1054]" gridrow="15" pageId="15" pageNumber="1203">
<th id="76A05DBE227A4FA024EDED45FEF2FBD2" box="[159,299,1033,1054]" gridcol="0" gridrow="15" pageId="15" pageNumber="1203">MOW</th>
<td id="76A05DBE227A4FA0253EED45FE5BFBD2" box="[332,386,1033,1054]" gridcol="1" gridrow="15" pageId="15" pageNumber="1203">80.6</td>
<td id="76A05DBE227A4FA025D1ED45FE00FBD2" box="[419,473,1033,1054]" gridcol="2" gridrow="15" pageId="15" pageNumber="1203">9.9</td>
<td id="76A05DBE227A4FA0258DED45FDECFBD2" box="[511,565,1033,1054]" gridcol="3" gridrow="15" pageId="15" pageNumber="1203">99.9</td>
<td id="76A05DBE227A4FA02621ED45FD4DFBD2" box="[595,660,1033,1054]" gridcol="4" gridrow="15" pageId="15" pageNumber="1203">13.3</td>
<td id="76A05DBE227A4FA026CBED45FD36FBD2" box="[697,751,1033,1054]" gridcol="5" gridrow="15" pageId="15" pageNumber="1203">95.5</td>
<td id="76A05DBE227A4FA02762ED45FC9FFBD2" box="[784,838,1033,1054]" gridcol="6" gridrow="15" pageId="15" pageNumber="1203">11.2</td>
<td id="76A05DBE227A4FA02715ED45FC4BFBD2" box="[871,914,1033,1054]" gridcol="7" gridrow="15" pageId="15" pageNumber="1203">13.6</td>
<td id="76A05DBE227A4FA027BBED45FC30FBD2" box="[969,1001,1033,1054]" gridcol="8" gridrow="15" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02061ED45FBE4FBD2" box="[1043,1085,1033,1054]" gridcol="9" gridrow="15" pageId="15" pageNumber="1203">12.7</td>
<td id="76A05DBE227A4FA02017ED45FB7BFBD2" box="[1125,1186,1033,1054]" gridcol="10" gridrow="15" pageId="15" pageNumber="1203">12.8</td>
<td id="76A05DBE227A4FA020AAED45FAC0FBD2" box="[1240,1305,1033,1054]" gridcol="11" gridrow="15" pageId="15" pageNumber="1203">87.1</td>
<td id="76A05DBE227A4FA0211EED45FA77FBD2" box="[1388,1454,1033,1054]" gridcol="12" gridrow="15" pageId="15" pageNumber="1203">13.9</td>
</tr>
<tr id="357134C2227A4FA024EDED65FA77FBF2" box="[159,1454,1065,1086]" gridrow="16" pageId="15" pageNumber="1203">
<th id="76A05DBE227A4FA024EDED65FEF2FBF2" box="[159,299,1065,1086]" gridcol="0" gridrow="16" pageId="15" pageNumber="1203">P2A</th>
<td id="76A05DBE227A4FA0253EED65FE5BFBF2" box="[332,386,1065,1086]" gridcol="1" gridrow="16" pageId="15" pageNumber="1203">77.2</td>
<td id="76A05DBE227A4FA025D1ED65FE00FBF2" box="[419,473,1065,1086]" gridcol="2" gridrow="16" pageId="15" pageNumber="1203">9.5</td>
<td id="76A05DBE227A4FA0258DED65FDECFBF2" box="[511,565,1065,1086]" gridcol="3" gridrow="16" pageId="15" pageNumber="1203">103.1</td>
<td id="76A05DBE227A4FA02621ED65FD4DFBF2" box="[595,660,1065,1086]" gridcol="4" gridrow="16" pageId="15" pageNumber="1203">13.7</td>
<td id="76A05DBE227A4FA026CBED65FD36FBF2" box="[697,751,1065,1086]" gridcol="5" gridrow="16" pageId="15" pageNumber="1203">88.7</td>
<td id="76A05DBE227A4FA02762ED65FC9FFBF2" box="[784,838,1065,1086]" gridcol="6" gridrow="16" pageId="15" pageNumber="1203">10.4</td>
<td id="76A05DBE227A4FA02715ED65FC4BFBF2" box="[871,914,1065,1086]" gridcol="7" gridrow="16" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA027BBED65FC30FBF2" box="[969,1001,1065,1086]" gridcol="8" gridrow="16" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02061ED65FBE4FBF2" box="[1043,1085,1065,1086]" gridcol="9" gridrow="16" pageId="15" pageNumber="1203">8.1</td>
<td id="76A05DBE227A4FA02017ED65FB7BFBF2" box="[1125,1186,1065,1086]" gridcol="10" gridrow="16" pageId="15" pageNumber="1203">8.2</td>
<td id="76A05DBE227A4FA020AAED65FAC0FBF2" box="[1240,1305,1065,1086]" gridcol="11" gridrow="16" pageId="15" pageNumber="1203">141.6</td>
<td id="76A05DBE227A4FA0211EED65FA77FBF2" box="[1388,1454,1065,1086]" gridcol="12" gridrow="16" pageId="15" pageNumber="1203">22.6</td>
</tr>
<tr id="357134C2227A4FA024EDED05FA77FB92" box="[159,1454,1097,1118]" gridrow="17" pageId="15" pageNumber="1203">
<th id="76A05DBE227A4FA024EDED05FEF2FB92" box="[159,299,1097,1118]" gridcol="0" gridrow="17" pageId="15" pageNumber="1203">P2B</th>
<td id="76A05DBE227A4FA0253EED05FE5BFB92" box="[332,386,1097,1118]" gridcol="1" gridrow="17" pageId="15" pageNumber="1203">134.6</td>
<td id="76A05DBE227A4FA025D1ED05FE00FB92" box="[419,473,1097,1118]" gridcol="2" gridrow="17" pageId="15" pageNumber="1203">16.5</td>
<td id="76A05DBE227A4FA0258DED05FDECFB92" box="[511,565,1097,1118]" gridcol="3" gridrow="17" pageId="15" pageNumber="1203">108.3</td>
<td id="76A05DBE227A4FA02621ED05FD4DFB92" box="[595,660,1097,1118]" gridcol="4" gridrow="17" pageId="15" pageNumber="1203">14.4</td>
<td id="76A05DBE227A4FA026CBED05FD36FB92" box="[697,751,1097,1118]" gridcol="5" gridrow="17" pageId="15" pageNumber="1203">81.3</td>
<td id="76A05DBE227A4FA02762ED05FC9FFB92" box="[784,838,1097,1118]" gridcol="6" gridrow="17" pageId="15" pageNumber="1203">9.6</td>
<td id="76A05DBE227A4FA02715ED05FC4BFB92" box="[871,914,1097,1118]" gridcol="7" gridrow="17" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA027BBED05FC30FB92" box="[969,1001,1097,1118]" gridcol="8" gridrow="17" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02061ED05FBE4FB92" box="[1043,1085,1097,1118]" gridcol="9" gridrow="17" pageId="15" pageNumber="1203">11.7</td>
<td id="76A05DBE227A4FA02017ED05FB7BFB92" box="[1125,1186,1097,1118]" gridcol="10" gridrow="17" pageId="15" pageNumber="1203">11.8</td>
<td id="76A05DBE227A4FA020AAED05FAC0FB92" box="[1240,1305,1097,1118]" gridcol="11" gridrow="17" pageId="15" pageNumber="1203">107.9</td>
<td id="76A05DBE227A4FA0211EED05FA77FB92" box="[1388,1454,1097,1118]" gridcol="12" gridrow="17" pageId="15" pageNumber="1203">17.2</td>
</tr>
<tr id="357134C2227A4FA024EDED24FA77FBB1" box="[159,1454,1128,1149]" gridrow="18" pageId="15" pageNumber="1203">
<th id="76A05DBE227A4FA024EDED24FEF2FBB1" box="[159,299,1128,1149]" gridcol="0" gridrow="18" pageId="15" pageNumber="1203">P2S</th>
<td id="76A05DBE227A4FA0253EED24FE5BFBB1" box="[332,386,1128,1149]" gridcol="1" gridrow="18" pageId="15" pageNumber="1203">206.8</td>
<td id="76A05DBE227A4FA025D1ED24FE00FBB1" box="[419,473,1128,1149]" gridcol="2" gridrow="18" pageId="15" pageNumber="1203">25.3</td>
<td id="76A05DBE227A4FA0258DED24FDECFBB1" box="[511,565,1128,1149]" gridcol="3" gridrow="18" pageId="15" pageNumber="1203">208.5</td>
<td id="76A05DBE227A4FA02621ED24FD4DFBB1" box="[595,660,1128,1149]" gridcol="4" gridrow="18" pageId="15" pageNumber="1203">27.8</td>
<td id="76A05DBE227A4FA026CBED24FD36FBB1" box="[697,751,1128,1149]" gridcol="5" gridrow="18" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02762ED24FC9FFBB1" box="[784,838,1128,1149]" gridcol="6" gridrow="18" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02715ED24FC4BFBB1" box="[871,914,1128,1149]" gridcol="7" gridrow="18" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA027BBED24FC30FBB1" box="[969,1001,1128,1149]" gridcol="8" gridrow="18" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02061ED24FBE4FBB1" box="[1043,1085,1128,1149]" gridcol="9" gridrow="18" pageId="15" pageNumber="1203">23.7</td>
<td id="76A05DBE227A4FA02017ED24FB7BFBB1" box="[1125,1186,1128,1149]" gridcol="10" gridrow="18" pageId="15" pageNumber="1203">23.9</td>
<td id="76A05DBE227A4FA020AAED24FAC0FBB1" box="[1240,1305,1128,1149]" gridcol="11" gridrow="18" pageId="15" pageNumber="1203">211.9</td>
<td id="76A05DBE227A4FA0211EED24FA77FBB1" box="[1388,1454,1128,1149]" gridcol="12" gridrow="18" pageId="15" pageNumber="1203">33.8</td>
</tr>
<tr id="357134C2227A4FA024EDEDC4FA77FB51" box="[159,1454,1160,1181]" gridrow="19" pageId="15" pageNumber="1203">
<th id="76A05DBE227A4FA024EDEDC4FEF2FB51" box="[159,299,1160,1181]" gridcol="0" gridrow="19" pageId="15" pageNumber="1203">PCS</th>
<td id="76A05DBE227A4FA0253EEDC4FE5BFB51" box="[332,386,1160,1181]" gridcol="1" gridrow="19" pageId="15" pageNumber="1203">356.0</td>
<td id="76A05DBE227A4FA025D1EDC4FE00FB51" box="[419,473,1160,1181]" gridcol="2" gridrow="19" pageId="15" pageNumber="1203">43.6</td>
<td id="76A05DBE227A4FA0258DEDC4FDECFB51" box="[511,565,1160,1181]" gridcol="3" gridrow="19" pageId="15" pageNumber="1203">358.5</td>
<td id="76A05DBE227A4FA02621EDC4FD4DFB51" box="[595,660,1160,1181]" gridcol="4" gridrow="19" pageId="15" pageNumber="1203">47.8</td>
<td id="76A05DBE227A4FA026CBEDC4FD36FB51" box="[697,751,1160,1181]" gridcol="5" gridrow="19" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02762EDC4FC9FFB51" box="[784,838,1160,1181]" gridcol="6" gridrow="19" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02715EDC4FC4BFB51" box="[871,914,1160,1181]" gridcol="7" gridrow="19" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA027BBEDC4FC30FB51" box="[969,1001,1160,1181]" gridcol="8" gridrow="19" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02061EDC4FBE4FB51" box="[1043,1085,1160,1181]" gridcol="9" gridrow="19" pageId="15" pageNumber="1203">44.9</td>
<td id="76A05DBE227A4FA02017EDC4FB7BFB51" box="[1125,1186,1160,1181]" gridcol="10" gridrow="19" pageId="15" pageNumber="1203">45.4</td>
<td id="76A05DBE227A4FA020AAEDC4FAC0FB51" box="[1240,1305,1160,1181]" gridcol="11" gridrow="19" pageId="15" pageNumber="1203">241.7</td>
<td id="76A05DBE227A4FA0211EEDC4FA77FB51" box="[1388,1454,1160,1181]" gridcol="12" gridrow="19" pageId="15" pageNumber="1203">38.6</td>
</tr>
<tr id="357134C2227A4FA024EDEDE4FA77FB71" box="[159,1454,1192,1213]" gridrow="20" pageId="15" pageNumber="1203">
<th id="76A05DBE227A4FA024EDEDE4FEF2FB71" box="[159,299,1192,1213]" gridcol="0" gridrow="20" pageId="15" pageNumber="1203">PD1</th>
<td id="76A05DBE227A4FA0253EEDE4FE5BFB71" box="[332,386,1192,1213]" gridcol="1" gridrow="20" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA025D1EDE4FE00FB71" box="[419,473,1192,1213]" gridcol="2" gridrow="20" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA0258DEDE4FDECFB71" box="[511,565,1192,1213]" gridcol="3" gridrow="20" pageId="15" pageNumber="1203">468.1</td>
<td id="76A05DBE227A4FA02621EDE4FD4DFB71" box="[595,660,1192,1213]" gridcol="4" gridrow="20" pageId="15" pageNumber="1203">62.4</td>
<td id="76A05DBE227A4FA026CBEDE4FD36FB71" box="[697,751,1192,1213]" gridcol="5" gridrow="20" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02762EDE4FC9FFB71" box="[784,838,1192,1213]" gridcol="6" gridrow="20" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02715EDE4FC4BFB71" box="[871,914,1192,1213]" gridcol="7" gridrow="20" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA027BBEDE4FC30FB71" box="[969,1001,1192,1213]" gridcol="8" gridrow="20" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02061EDE4FBE4FB71" box="[1043,1085,1192,1213]" gridcol="9" gridrow="20" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02017EDE4FB7BFB71" box="[1125,1186,1192,1213]" gridcol="10" gridrow="20" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA020AAEDE4FAC0FB71" box="[1240,1305,1192,1213]" gridcol="11" gridrow="20" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA0211EEDE4FA77FB71" box="[1388,1454,1192,1213]" gridcol="12" gridrow="20" pageId="15" pageNumber="1203"></td>
</tr>
<tr id="357134C2227A4FA024EDED84FA77FB11" box="[159,1454,1224,1245]" gridrow="21" pageId="15" pageNumber="1203">
<th id="76A05DBE227A4FA024EDED84FEF2FB11" box="[159,299,1224,1245]" gridcol="0" gridrow="21" pageId="15" pageNumber="1203">PD2</th>
<td id="76A05DBE227A4FA0253EED84FE5BFB11" box="[332,386,1224,1245]" gridcol="1" gridrow="21" pageId="15" pageNumber="1203">632.3</td>
<td id="76A05DBE227A4FA025D1ED84FE00FB11" box="[419,473,1224,1245]" gridcol="2" gridrow="21" pageId="15" pageNumber="1203">77.4</td>
<td id="76A05DBE227A4FA0258DED84FDECFB11" box="[511,565,1224,1245]" gridcol="3" gridrow="21" pageId="15" pageNumber="1203">586.3</td>
<td id="76A05DBE227A4FA02621ED84FD4DFB11" box="[595,660,1224,1245]" gridcol="4" gridrow="21" pageId="15" pageNumber="1203">78.2</td>
<td id="76A05DBE227A4FA026CBED84FD36FB11" box="[697,751,1224,1245]" gridcol="5" gridrow="21" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02762ED84FC9FFB11" box="[784,838,1224,1245]" gridcol="6" gridrow="21" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02715ED84FC4BFB11" box="[871,914,1224,1245]" gridcol="7" gridrow="21" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA027BBED84FC30FB11" box="[969,1001,1224,1245]" gridcol="8" gridrow="21" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02061ED84FBE4FB11" box="[1043,1085,1224,1245]" gridcol="9" gridrow="21" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02017ED84FB7BFB11" box="[1125,1186,1224,1245]" gridcol="10" gridrow="21" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA020AAED84FAC0FB11" box="[1240,1305,1224,1245]" gridcol="11" gridrow="21" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA0211EED84FA77FB11" box="[1388,1454,1224,1245]" gridcol="12" gridrow="21" pageId="15" pageNumber="1203"></td>
</tr>
<tr id="357134C2227A4FA024EDEDA4FA77FB31" box="[159,1454,1256,1277]" gridrow="22" pageId="15" pageNumber="1203">
<th id="76A05DBE227A4FA024EDEDA4FEF2FB31" box="[159,299,1256,1277]" gridcol="0" gridrow="22" pageId="15" pageNumber="1203">PDI</th>
<td id="76A05DBE227A4FA0253EEDA4FE5BFB31" box="[332,386,1256,1277]" gridcol="1" gridrow="22" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA025D1EDA4FE00FB31" box="[419,473,1256,1277]" gridcol="2" gridrow="22" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA0258DEDA4FDECFB31" box="[511,565,1256,1277]" gridcol="3" gridrow="22" pageId="15" pageNumber="1203">83.1</td>
<td id="76A05DBE227A4FA02621EDA4FD4DFB31" box="[595,660,1256,1277]" gridcol="4" gridrow="22" pageId="15" pageNumber="1203">11.1</td>
<td id="76A05DBE227A4FA026CBEDA4FD36FB31" box="[697,751,1256,1277]" gridcol="5" gridrow="22" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02762EDA4FC9FFB31" box="[784,838,1256,1277]" gridcol="6" gridrow="22" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02715EDA4FC4BFB31" box="[871,914,1256,1277]" gridcol="7" gridrow="22" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA027BBEDA4FC30FB31" box="[969,1001,1256,1277]" gridcol="8" gridrow="22" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02061EDA4FBE4FB31" box="[1043,1085,1256,1277]" gridcol="9" gridrow="22" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02017EDA4FB7BFB31" box="[1125,1186,1256,1277]" gridcol="10" gridrow="22" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA020AAEDA4FAC0FB31" box="[1240,1305,1256,1277]" gridcol="11" gridrow="22" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA0211EEDA4FA77FB31" box="[1388,1454,1256,1277]" gridcol="12" gridrow="22" pageId="15" pageNumber="1203"></td>
</tr>
<tr id="357134C2227A4FA024EDEC44FA77FAD1" box="[159,1454,1288,1309]" gridrow="23" pageId="15" pageNumber="1203">
<th id="76A05DBE227A4FA024EDEC44FEF2FAD1" box="[159,299,1288,1309]" gridcol="0" gridrow="23" pageId="15" pageNumber="1203">PDO</th>
<td id="76A05DBE227A4FA0253EEC44FE5BFAD1" box="[332,386,1288,1309]" gridcol="1" gridrow="23" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA025D1EC44FE00FAD1" box="[419,473,1288,1309]" gridcol="2" gridrow="23" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA0258DEC44FDECFAD1" box="[511,565,1288,1309]" gridcol="3" gridrow="23" pageId="15" pageNumber="1203">181.3</td>
<td id="76A05DBE227A4FA02621EC44FD4DFAD1" box="[595,660,1288,1309]" gridcol="4" gridrow="23" pageId="15" pageNumber="1203">24.2</td>
<td id="76A05DBE227A4FA026CBEC44FD36FAD1" box="[697,751,1288,1309]" gridcol="5" gridrow="23" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02762EC44FC9FFAD1" box="[784,838,1288,1309]" gridcol="6" gridrow="23" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02715EC44FC4BFAD1" box="[871,914,1288,1309]" gridcol="7" gridrow="23" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA027BBEC44FC30FAD1" box="[969,1001,1288,1309]" gridcol="8" gridrow="23" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02061EC44FBE4FAD1" box="[1043,1085,1288,1309]" gridcol="9" gridrow="23" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02017EC44FB7BFAD1" box="[1125,1186,1288,1309]" gridcol="10" gridrow="23" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA020AAEC44FAC0FAD1" box="[1240,1305,1288,1309]" gridcol="11" gridrow="23" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA0211EEC44FA77FAD1" box="[1388,1454,1288,1309]" gridcol="12" gridrow="23" pageId="15" pageNumber="1203"></td>
</tr>
<tr id="357134C2227A4FA024EDEC64FA77FAF1" box="[159,1454,1320,1341]" gridrow="24" pageId="15" pageNumber="1203">
<th id="76A05DBE227A4FA024EDEC64FEF2FAF1" box="[159,299,1320,1341]" gridcol="0" gridrow="24" pageId="15" pageNumber="1203">PGW</th>
<td id="76A05DBE227A4FA0253EEC64FE5BFAF1" box="[332,386,1320,1341]" gridcol="1" gridrow="24" pageId="15" pageNumber="1203">198.4</td>
<td id="76A05DBE227A4FA025D1EC64FE00FAF1" box="[419,473,1320,1341]" gridcol="2" gridrow="24" pageId="15" pageNumber="1203">24.3</td>
<td id="76A05DBE227A4FA0258DEC64FDECFAF1" box="[511,565,1320,1341]" gridcol="3" gridrow="24" pageId="15" pageNumber="1203">187.2</td>
<td id="76A05DBE227A4FA02621EC64FD4DFAF1" box="[595,660,1320,1341]" gridcol="4" gridrow="24" pageId="15" pageNumber="1203">25.0</td>
<td id="76A05DBE227A4FA026CBEC64FD36FAF1" box="[697,751,1320,1341]" gridcol="5" gridrow="24" pageId="15" pageNumber="1203">195.2</td>
<td id="76A05DBE227A4FA02762EC64FC9FFAF1" box="[784,838,1320,1341]" gridcol="6" gridrow="24" pageId="15" pageNumber="1203">23.0</td>
<td id="76A05DBE227A4FA02715EC64FC4BFAF1" box="[871,914,1320,1341]" gridcol="7" gridrow="24" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA027BBEC64FC30FAF1" box="[969,1001,1320,1341]" gridcol="8" gridrow="24" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02061EC64FBE4FAF1" box="[1043,1085,1320,1341]" gridcol="9" gridrow="24" pageId="15" pageNumber="1203">23.7</td>
<td id="76A05DBE227A4FA02017EC64FB7BFAF1" box="[1125,1186,1320,1341]" gridcol="10" gridrow="24" pageId="15" pageNumber="1203">23.9</td>
<td id="76A05DBE227A4FA020AAEC64FAC0FAF1" box="[1240,1305,1320,1341]" gridcol="11" gridrow="24" pageId="15" pageNumber="1203">172.2</td>
<td id="76A05DBE227A4FA0211EEC64FA77FAF1" box="[1388,1454,1320,1341]" gridcol="12" gridrow="24" pageId="15" pageNumber="1203">27.5</td>
</tr>
<tr id="357134C2227A4FA024EDEC04FA77FA91" box="[159,1454,1352,1373]" gridrow="25" pageId="15" pageNumber="1203">
<th id="76A05DBE227A4FA024EDEC04FEF2FA91" box="[159,299,1352,1373]" gridcol="0" gridrow="25" pageId="15" pageNumber="1203">PIW</th>
<td id="76A05DBE227A4FA0253EEC04FE5BFA91" box="[332,386,1352,1373]" gridcol="1" gridrow="25" pageId="15" pageNumber="1203">150.3</td>
<td id="76A05DBE227A4FA025D1EC04FE00FA91" box="[419,473,1352,1373]" gridcol="2" gridrow="25" pageId="15" pageNumber="1203">18.4</td>
<td id="76A05DBE227A4FA0258DEC04FDECFA91" box="[511,565,1352,1373]" gridcol="3" gridrow="25" pageId="15" pageNumber="1203">173.7</td>
<td id="76A05DBE227A4FA02621EC04FD4DFA91" box="[595,660,1352,1373]" gridcol="4" gridrow="25" pageId="15" pageNumber="1203">23.2</td>
<td id="76A05DBE227A4FA026CBEC04FD36FA91" box="[697,751,1352,1373]" gridcol="5" gridrow="25" pageId="15" pageNumber="1203">180.3</td>
<td id="76A05DBE227A4FA02762EC04FC9FFA91" box="[784,838,1352,1373]" gridcol="6" gridrow="25" pageId="15" pageNumber="1203">21.2</td>
<td id="76A05DBE227A4FA02715EC04FC4BFA91" box="[871,914,1352,1373]" gridcol="7" gridrow="25" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA027BBEC04FC30FA91" box="[969,1001,1352,1373]" gridcol="8" gridrow="25" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02061EC04FBE4FA91" box="[1043,1085,1352,1373]" gridcol="9" gridrow="25" pageId="15" pageNumber="1203">22.1</td>
<td id="76A05DBE227A4FA02017EC04FB7BFA91" box="[1125,1186,1352,1373]" gridcol="10" gridrow="25" pageId="15" pageNumber="1203">22.3</td>
<td id="76A05DBE227A4FA020AAEC04FAC0FA91" box="[1240,1305,1352,1373]" gridcol="11" gridrow="25" pageId="15" pageNumber="1203">173.0</td>
<td id="76A05DBE227A4FA0211EEC04FA77FA91" box="[1388,1454,1352,1373]" gridcol="12" gridrow="25" pageId="15" pageNumber="1203">27.6</td>
</tr>
<tr id="357134C2227A4FA024EDEC2BFA77FAB0" box="[159,1454,1383,1404]" gridrow="26" pageId="15" pageNumber="1203">
<th id="76A05DBE227A4FA024EDEC2BFEF2FAB0" box="[159,299,1383,1404]" gridcol="0" gridrow="26" pageId="15" pageNumber="1203">POR</th>
<td id="76A05DBE227A4FA0253EEC2BFE5BFAB0" box="[332,386,1383,1404]" gridcol="1" gridrow="26" pageId="15" pageNumber="1203">73.0</td>
<td id="76A05DBE227A4FA025D1EC2BFE00FAB0" box="[419,473,1383,1404]" gridcol="2" gridrow="26" pageId="15" pageNumber="1203">8.9</td>
<td id="76A05DBE227A4FA0258DEC2BFDECFAB0" box="[511,565,1383,1404]" gridcol="3" gridrow="26" pageId="15" pageNumber="1203">62.6</td>
<td id="76A05DBE227A4FA02621EC2BFD4DFAB0" box="[595,660,1383,1404]" gridcol="4" gridrow="26" pageId="15" pageNumber="1203">8.3</td>
<td id="76A05DBE227A4FA026CBEC2BFD36FAB0" box="[697,751,1383,1404]" gridcol="5" gridrow="26" pageId="15" pageNumber="1203">85.4</td>
<td id="76A05DBE227A4FA02762EC2BFC9FFAB0" box="[784,838,1383,1404]" gridcol="6" gridrow="26" pageId="15" pageNumber="1203">10.0</td>
<td id="76A05DBE227A4FA02715EC2BFC4BFAB0" box="[871,914,1383,1404]" gridcol="7" gridrow="26" pageId="15" pageNumber="1203">7.7</td>
<td id="76A05DBE227A4FA027BBEC2BFC30FAB0" box="[969,1001,1383,1404]" gridcol="8" gridrow="26" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02061EC2BFBE4FAB0" box="[1043,1085,1383,1404]" gridcol="9" gridrow="26" pageId="15" pageNumber="1203">9.6</td>
<td id="76A05DBE227A4FA02017EC2BFB7BFAB0" box="[1125,1186,1383,1404]" gridcol="10" gridrow="26" pageId="15" pageNumber="1203">9.7</td>
<td id="76A05DBE227A4FA020AAEC2BFAC0FAB0" box="[1240,1305,1383,1404]" gridcol="11" gridrow="26" pageId="15" pageNumber="1203">56.2</td>
<td id="76A05DBE227A4FA0211EEC2BFA77FAB0" box="[1388,1454,1383,1404]" gridcol="12" gridrow="26" pageId="15" pageNumber="1203">9.0</td>
</tr>
<tr id="357134C2227A4FA024EDECCBFA77FA50" box="[159,1454,1415,1436]" gridrow="27" pageId="15" pageNumber="1203">
<th id="76A05DBE227A4FA024EDECCBFEF2FA50" box="[159,299,1415,1436]" gridcol="0" gridrow="27" pageId="15" pageNumber="1203">PP2</th>
<td id="76A05DBE227A4FA0253EECCBFE5BFA50" box="[332,386,1415,1436]" gridcol="1" gridrow="27" pageId="15" pageNumber="1203">328.5</td>
<td id="76A05DBE227A4FA025D1ECCBFE00FA50" box="[419,473,1415,1436]" gridcol="2" gridrow="27" pageId="15" pageNumber="1203">40.2</td>
<td id="76A05DBE227A4FA0258DECCBFDECFA50" box="[511,565,1415,1436]" gridcol="3" gridrow="27" pageId="15" pageNumber="1203">307.3</td>
<td id="76A05DBE227A4FA02621ECCBFD4DFA50" box="[595,660,1415,1436]" gridcol="4" gridrow="27" pageId="15" pageNumber="1203">41.0</td>
<td id="76A05DBE227A4FA026CBECCBFD36FA50" box="[697,751,1415,1436]" gridcol="5" gridrow="27" pageId="15" pageNumber="1203">348.4</td>
<td id="76A05DBE227A4FA02762ECCBFC9FFA50" box="[784,838,1415,1436]" gridcol="6" gridrow="27" pageId="15" pageNumber="1203">41.0</td>
<td id="76A05DBE227A4FA02715ECCBFC4BFA50" box="[871,914,1415,1436]" gridcol="7" gridrow="27" pageId="15" pageNumber="1203">34.8</td>
<td id="76A05DBE227A4FA027BBECCBFC30FA50" box="[969,1001,1415,1436]" gridcol="8" gridrow="27" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02061ECCBFBE4FA50" box="[1043,1085,1415,1436]" gridcol="9" gridrow="27" pageId="15" pageNumber="1203">44.4</td>
<td id="76A05DBE227A4FA02017ECCBFB7BFA50" box="[1125,1186,1415,1436]" gridcol="10" gridrow="27" pageId="15" pageNumber="1203">44.8</td>
<td id="76A05DBE227A4FA020AAECCBFAC0FA50" box="[1240,1305,1415,1436]" gridcol="11" gridrow="27" pageId="15" pageNumber="1203">299.0</td>
<td id="76A05DBE227A4FA0211EECCBFA77FA50" box="[1388,1454,1415,1436]" gridcol="12" gridrow="27" pageId="15" pageNumber="1203">47.7</td>
</tr>
<tr id="357134C2227A4FA024EDECEBFA77FA70" box="[159,1454,1447,1468]" gridrow="28" pageId="15" pageNumber="1203">
<th id="76A05DBE227A4FA024EDECEBFEF2FA70" box="[159,299,1447,1468]" gridcol="0" gridrow="28" pageId="15" pageNumber="1203">SL</th>
<td id="76A05DBE227A4FA0253EECEBFE5BFA70" box="[332,386,1447,1468]" gridcol="1" gridrow="28" pageId="15" pageNumber="1203">816.8</td>
<td id="76A05DBE227A4FA025D1ECEBFE00FA70" box="[419,473,1447,1468]" gridcol="2" gridrow="28" pageId="15" pageNumber="1203">100.0</td>
<td id="76A05DBE227A4FA0258DECEBFDECFA70" box="[511,565,1447,1468]" gridcol="3" gridrow="28" pageId="15" pageNumber="1203">750.2</td>
<td id="76A05DBE227A4FA02621ECEBFD4DFA70" box="[595,660,1447,1468]" gridcol="4" gridrow="28" pageId="15" pageNumber="1203">100.0</td>
<td id="76A05DBE227A4FA026CBECEBFD36FA70" box="[697,751,1447,1468]" gridcol="5" gridrow="28" pageId="15" pageNumber="1203">850.4</td>
<td id="76A05DBE227A4FA02762ECEBFC9FFA70" box="[784,838,1447,1468]" gridcol="6" gridrow="28" pageId="15" pageNumber="1203">100.0</td>
<td id="76A05DBE227A4FA02715ECEBFC4BFA70" box="[871,914,1447,1468]" gridcol="7" gridrow="28" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA027BBECEBFC30FA70" box="[969,1001,1447,1468]" gridcol="8" gridrow="28" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02061ECEBFBE4FA70" box="[1043,1085,1447,1468]" gridcol="9" gridrow="28" pageId="15" pageNumber="1203">99.0</td>
<td id="76A05DBE227A4FA02017ECEBFB7BFA70" box="[1125,1186,1447,1468]" gridcol="10" gridrow="28" pageId="15" pageNumber="1203">100.0</td>
<td id="76A05DBE227A4FA020AAECEBFAC0FA70" box="[1240,1305,1447,1468]" gridcol="11" gridrow="28" pageId="15" pageNumber="1203">626.2</td>
<td id="76A05DBE227A4FA0211EECEBFA77FA70" box="[1388,1454,1447,1468]" gridcol="12" gridrow="28" pageId="15" pageNumber="1203">100.0</td>
</tr>
<tr id="357134C2227A4FA024EDEC8BFA77FA10" box="[159,1454,1479,1500]" gridrow="29" pageId="15" pageNumber="1203">
<th id="76A05DBE227A4FA024EDEC8BFEF2FA10" box="[159,299,1479,1500]" gridcol="0" gridrow="29" pageId="15" pageNumber="1203">TAL</th>
<td id="76A05DBE227A4FA0253EEC8BFE5BFA10" box="[332,386,1479,1500]" gridcol="1" gridrow="29" pageId="15" pageNumber="1203">493.5</td>
<td id="76A05DBE227A4FA025D1EC8BFE00FA10" box="[419,473,1479,1500]" gridcol="2" gridrow="29" pageId="15" pageNumber="1203">60.4</td>
<td id="76A05DBE227A4FA0258DEC8BFDECFA10" box="[511,565,1479,1500]" gridcol="3" gridrow="29" pageId="15" pageNumber="1203">436.8</td>
<td id="76A05DBE227A4FA02621EC8BFD4DFA10" box="[595,660,1479,1500]" gridcol="4" gridrow="29" pageId="15" pageNumber="1203">58.2</td>
<td id="76A05DBE227A4FA026CBEC8BFD36FA10" box="[697,751,1479,1500]" gridcol="5" gridrow="29" pageId="15" pageNumber="1203">502.0</td>
<td id="76A05DBE227A4FA02762EC8BFC9FFA10" box="[784,838,1479,1500]" gridcol="6" gridrow="29" pageId="15" pageNumber="1203">59.0</td>
<td id="76A05DBE227A4FA02715EC8BFC4BFA10" box="[871,914,1479,1500]" gridcol="7" gridrow="29" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA027BBEC8BFC30FA10" box="[969,1001,1479,1500]" gridcol="8" gridrow="29" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02061EC8BFBE4FA10" box="[1043,1085,1479,1500]" gridcol="9" gridrow="29" pageId="15" pageNumber="1203">55.4</td>
<td id="76A05DBE227A4FA02017EC8BFB7BFA10" box="[1125,1186,1479,1500]" gridcol="10" gridrow="29" pageId="15" pageNumber="1203">56.0</td>
<td id="76A05DBE227A4FA020AAEC8BFAC0FA10" box="[1240,1305,1479,1500]" gridcol="11" gridrow="29" pageId="15" pageNumber="1203">318.3</td>
<td id="76A05DBE227A4FA0211EEC8BFA77FA10" box="[1388,1454,1479,1500]" gridcol="12" gridrow="29" pageId="15" pageNumber="1203">50.8</td>
</tr>
<tr id="357134C2227A4FA024EDECABFA77FA30" box="[159,1454,1511,1532]" gridrow="30" pageId="15" pageNumber="1203">
<th id="76A05DBE227A4FA024EDECABFEF2FA30" box="[159,299,1511,1532]" gridcol="0" gridrow="30" pageId="15" pageNumber="1203">TBW</th>
<td id="76A05DBE227A4FA0253EECABFE5BFA30" box="[332,386,1511,1532]" gridcol="1" gridrow="30" pageId="15" pageNumber="1203">94.8</td>
<td id="76A05DBE227A4FA025D1ECABFE00FA30" box="[419,473,1511,1532]" gridcol="2" gridrow="30" pageId="15" pageNumber="1203">11.6</td>
<td id="76A05DBE227A4FA0258DECABFDECFA30" box="[511,565,1511,1532]" gridcol="3" gridrow="30" pageId="15" pageNumber="1203">109.7</td>
<td id="76A05DBE227A4FA02621ECABFD4DFA30" box="[595,660,1511,1532]" gridcol="4" gridrow="30" pageId="15" pageNumber="1203">14.6</td>
<td id="76A05DBE227A4FA026CBECABFD36FA30" box="[697,751,1511,1532]" gridcol="5" gridrow="30" pageId="15" pageNumber="1203">132.7</td>
<td id="76A05DBE227A4FA02762ECABFC9FFA30" box="[784,838,1511,1532]" gridcol="6" gridrow="30" pageId="15" pageNumber="1203">15.6</td>
<td id="76A05DBE227A4FA02715ECABFC4BFA30" box="[871,914,1511,1532]" gridcol="7" gridrow="30" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA027BBECABFC30FA30" box="[969,1001,1511,1532]" gridcol="8" gridrow="30" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02061ECABFBE4FA30" box="[1043,1085,1511,1532]" gridcol="9" gridrow="30" pageId="15" pageNumber="1203">13.2</td>
<td id="76A05DBE227A4FA02017ECABFB7BFA30" box="[1125,1186,1511,1532]" gridcol="10" gridrow="30" pageId="15" pageNumber="1203">13.3</td>
<td id="76A05DBE227A4FA020AAECABFAC0FA30" box="[1240,1305,1511,1532]" gridcol="11" gridrow="30" pageId="15" pageNumber="1203">103.5</td>
<td id="76A05DBE227A4FA0211EECABFA77FA30" box="[1388,1454,1511,1532]" gridcol="12" gridrow="30" pageId="15" pageNumber="1203">16.5</td>
</tr>
<tr id="357134C2227A4FA024EDEF4BFA77F9D0" box="[159,1454,1543,1564]" gridrow="31" pageId="15" pageNumber="1203">
<th id="76A05DBE227A4FA024EDEF4BFEF2F9D0" box="[159,299,1543,1564]" gridcol="0" gridrow="31" pageId="15" pageNumber="1203">TL</th>
<td id="76A05DBE227A4FA0253EEF4BFE5BF9D0" box="[332,386,1543,1564]" gridcol="1" gridrow="31" pageId="15" pageNumber="1203">897.7</td>
<td id="76A05DBE227A4FA025D1EF4BFE00F9D0" box="[419,473,1543,1564]" gridcol="2" gridrow="31" pageId="15" pageNumber="1203">109.9</td>
<td id="76A05DBE227A4FA0258DEF4BFDECF9D0" box="[511,565,1543,1564]" gridcol="3" gridrow="31" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02621EF4BFD4DF9D0" box="[595,660,1543,1564]" gridcol="4" gridrow="31" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA026CBEF4BFD36F9D0" box="[697,751,1543,1564]" gridcol="5" gridrow="31" pageId="15" pageNumber="1203">925.4</td>
<td id="76A05DBE227A4FA02762EF4BFC9FF9D0" box="[784,838,1543,1564]" gridcol="6" gridrow="31" pageId="15" pageNumber="1203">108.8</td>
<td id="76A05DBE227A4FA02715EF4BFC4BF9D0" box="[871,914,1543,1564]" gridcol="7" gridrow="31" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA027BBEF4BFC30F9D0" box="[969,1001,1543,1564]" gridcol="8" gridrow="31" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02061EF4BFBE4F9D0" box="[1043,1085,1543,1564]" gridcol="9" gridrow="31" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA02017EF4BFB7BF9D0" box="[1125,1186,1543,1564]" gridcol="10" gridrow="31" pageId="15" pageNumber="1203"></td>
<td id="76A05DBE227A4FA020AAEF4BFAC0F9D0" box="[1240,1305,1543,1564]" gridcol="11" gridrow="31" pageId="15" pageNumber="1203">722.6</td>
<td id="76A05DBE227A4FA0211EEF4BFA77F9D0" box="[1388,1454,1543,1564]" gridcol="12" gridrow="31" pageId="15" pageNumber="1203">115.4</td>
</tr>
</table>
</paragraph>
<caption id="DF3E66082265B04F24EDE9F6FDA2FED3" pageId="16" pageNumber="1204" startId="16.[159,217,186,207]" targetBox="[159,1466,315,737]" targetIsTable="true" targetPageId="16" targetType="table">
<paragraph id="8BFE36802265B04F24EDE9F6FDA2FED3" blockId="16.[159,1467,186,287]" pageId="16" pageNumber="1204">
<emphasis id="B935EA922265B04F24EDE9F6FF28FF03" bold="true" box="[159,241,186,207]" pageId="16" pageNumber="1204">Table 2.</emphasis>
Summary of selected meristic features used to discriminate fossil and living genera of the family
<taxonomicName id="4C414D032265B04F20E8E9F6FADFFF03" authorityName="Gill" authorityYear="1862" box="[1178,1286,186,207]" class="Elasmobranchii" family="Narcinidae" kingdom="Animalia" order="Torpediniformes" pageId="16" pageNumber="1204" phylum="Chordata" rank="family">Narcinidae</taxonomicName>
. Includes new data from the examined material and data from
<bibRefCitation id="EFD04B712265B04F264FE998FCA8FF25" author="Fechhelm, J. D. &amp; McEachran, J. D." box="[573,881,212,233]" pageId="16" pageNumber="1204" pagination="171 - 209" refId="ref21620" refString="Fechhelm, J. D. &amp; McEachran, J. D. 1984. A revision of the electric ray genus Diplobatis with notes on the interrelationships of Narcinidae (Chondrichthyes, Torpediniformes). Bulletin of the Florida State Museum Biological Sciences, 29, 171 - 209." type="journal article" year="1984">Fechhelm &amp; McEachran (1984)</bibRefCitation>
, Rincon (1997),
<bibRefCitation id="EFD04B712265B04F2066E998FB38FF25" author="Carvalho, M. R. de &amp; Compagno, L. J. V. &amp; Last, P. R." box="[1044,1249,212,233]" pageId="16" pageNumber="1204" refId="ref20198" refString="Carvalho, M. R. de, Compagno, L. J. V. &amp; Last, P. R. 1999. Torpediniformes: Narcinidae, Numbfishes. Pp. 1397 - 2068 in K. E. Carpenter &amp; V. H. Niem (eds) FAO species identification guide for fishery purposes. The living marine resources of the Western Central Pacific. Volume 3. Batoid fishes, chimaeras and bony fishes part 1 (Elopidae to Linophrynidae). FAO, Rome." type="book" year="1999">
Carvalho
<emphasis id="B935EA922265B04F2000E998FB44FF25" box="[1138,1181,212,233]" italics="true" pageId="16" pageNumber="1204">et al</emphasis>
. (1999
</bibRefCitation>
,
<bibRefCitation id="EFD04B712265B04F209FE998FAFEFF25" author="Carvalho, M. R. de &amp; Compagno, L. J. V. &amp; Mee, J. K. L." box="[1261,1319,212,233]" pageId="16" pageNumber="1204" pagination="137 - 145" refId="ref20289" refString="Carvalho, M. R. de, Compagno, L. J. V. &amp; Mee, J. K. L. 2002 a. Narcine oculifera: a new species of electric ray from the gulfs of Oman and Aden (Chondrichthyes: Torpediniformes: Narcinidae). Copeia, 2002, 137 - 145." type="journal article" year="2002">2002a</bibRefCitation>
, b, 2003),
<bibRefCitation id="EFD04B712265B04F21E2E998FECAFEC8" author="Carvalho, M. R. de" pageId="16" pageNumber="1204" pagination="561 - 573" refId="ref19791" refString="Carvalho, M. R. de. 2001. A new species of electric ray, Narcine leoparda, from the tropical eastern Pacific Ocean (Chondrichthyes: Torpediniformes: Narcinidae). Proceedings of the Biological Society of Washington, 114 (3), 561 - 573." type="journal article" year="2001">Carvalho (2001</bibRefCitation>
,
<bibRefCitation id="EFD04B712265B04F256DE9A3FE96FEC8" author="Carvalho, M. R. de" box="[287,335,239,260]" pageId="16" pageNumber="1204" refId="ref19842" refString="Carvalho, M. R. de. 2008. New species of numbfishes from Australia, with a key to Australian electric rays of the genus Narcine Henle, 1834 (Chondrichthyes: Torpediniformes: Narcinidae). Pp. 241 - 260 in P. R. Last, W. T. White &amp; J. J. Pogonoski (eds) Description of new Australian chondrichthyans. CSIRO Marine and Atmospheric Research, Hobart." type="book" year="2008">2008</bibRefCitation>
), Rincon
<emphasis id="B935EA922265B04F25DDE9A3FE00FEC8" box="[431,473,239,260]" italics="true" pageId="16" pageNumber="1204">et al</emphasis>
. (2001), Carvalho &amp; Śeret (2002),
<bibRefCitation id="EFD04B712265B04F275EE9A3FBEFFEC8" author="Carvalho, M. R. de &amp; Randall, J. E." box="[812,1078,239,260]" pageId="16" pageNumber="1204" pagination="59 - 66" refId="ref19999" refString="Carvalho, M. R. de &amp; Randall, J. E. 2003. Numbfishes from the Arabian Sea and surrounding gulfs, with the description of a new species from Oman (Chondrichthyes: Torpediniformes: Narcinidae). Ichthyological Research, 50, 59 - 66." type="journal article" year="2003">Carvalho &amp; Randall (2003)</bibRefCitation>
, Menni
<emphasis id="B935EA922265B04F20F4E9A3FB69FEC8" box="[1158,1200,239,260]" italics="true" pageId="16" pageNumber="1204">et al</emphasis>
. (2008),
<bibRefCitation id="EFD04B712265B04F217AE9A3FF39FED3" author="Carvalho, M. R. de &amp; White, W. T." pageId="16" pageNumber="1204" pagination="149 - 160" refId="ref20097" refString="Carvalho, M. R. de &amp; White, W. T. 2016. Narcine baliensis, a new species of electric ray from southeast Asia (Chondrichthyes: Torpediniformes). Zootaxa, 4127, 149 - 160." type="journal article" year="2016">Carvalho &amp; White (2016)</bibRefCitation>
,
<bibRefCitation id="EFD04B712265B04F2499E846FE49FED3" author="Last, P. &amp; White, W. &amp; Carvalho, M. R. de &amp; Stehmann, M. &amp; Naylor, G." box="[235,400,266,287]" pageId="16" pageNumber="1204" refId="ref23787" refString="Last, P., White, W., Carvalho, M. R. de, S´eret, B., Stehmann, M. &amp; Naylor, G. 2016. Rays of the world. CSIRO Publishing, Clayton North, 800 pp." type="book" year="2016">
Last
<emphasis id="B935EA922265B04F2568E846FE9DFED3" box="[282,324,266,287]" italics="true" pageId="16" pageNumber="1204">et al</emphasis>
. (2016)
</bibRefCitation>
,
<bibRefCitation id="EFD04B712265B04F25E9E846FDAEFED3" author="Froese, R. &amp; Pauly, D." box="[411,631,266,287]" pageId="16" pageNumber="1204" refId="ref21888" refString="Froese, R. &amp; Pauly, D. 2017. FishBase. World Wide Web electronic publication version (02 / 2017). Updated at: http: // www. fishbase. org, accessed 13 April 2017." type="url" year="2017">Froese &amp; Pauly (2017)</bibRefCitation>
.
</paragraph>
</caption>
<paragraph id="8BFE36802265B04F24EDE870FA41FD2D" pageId="16" pageNumber="1204">
<table id="F941C42022654FA024EDE877FA63FD2D" box="[159,1466,315,737]" gridcols="6" gridrows="13" pageId="16" pageNumber="1204">
<tr id="357134C222654FA024EDE877FA63FE9D" box="[159,1466,315,337]" gridrow="0" pageId="16" pageNumber="1204">
<th id="76A05DBE22654FA024EDE877FE52FE9D" box="[159,395,315,337]" gridcol="0" gridrow="0" pageId="16" pageNumber="1204">Feature</th>
<th id="76A05DBE22654FA0258AE877FDB2FE9D" box="[504,619,315,337]" gridcol="1" gridrow="0" pageId="16" pageNumber="1204">
<taxonomicName id="4C414D032265B04F258AE870FDB2FE9D" authorityName="Alcock" authorityYear="1898" box="[504,619,316,337]" class="Elasmobranchii" family="Narcinidae" genus="Benthobatis" kingdom="Animalia" order="Torpediniformes" pageId="16" pageNumber="1204" phylum="Chordata" rank="genus">
<emphasis id="B935EA922265B04F258AE870FDB2FE9D" box="[504,619,316,337]" italics="true" pageId="16" pageNumber="1204">Benthobatis</emphasis>
</taxonomicName>
</th>
<th id="76A05DBE22654FA026A5E877FCE5FE9D" box="[727,828,315,337]" gridcol="2" gridrow="0" pageId="16" pageNumber="1204">
<taxonomicName id="4C414D032265B04F26A5E870FCE5FE9D" baseAuthorityName="Fechhelm &amp; McEachran" baseAuthorityYear="1984" box="[727,828,316,337]" class="Elasmobranchii" family="Narcinidae" genus="Diplobatis" kingdom="Animalia" order="Torpediniformes" pageId="16" pageNumber="1204" phylum="Chordata" rank="genus">
<emphasis id="B935EA922265B04F26A5E870FCE5FE9D" box="[727,828,316,337]" italics="true" pageId="16" pageNumber="1204">Diplobatis</emphasis>
</taxonomicName>
</th>
<th id="76A05DBE22654FA027DBE877FBD7FE9D" box="[937,1038,315,337]" gridcol="3" gridrow="0" pageId="16" pageNumber="1204">
<taxonomicName id="4C414D032265B04F27DBE870FBD7FE9D" authorityName="Heckel" authorityYear="1846" box="[937,1038,316,337]" class="Elasmobranchii" family="Narcinidae" genus="Discopyge" kingdom="Animalia" order="Torpediniformes" pageId="16" pageNumber="1204" phylum="Chordata" rank="genus">
<emphasis id="B935EA922265B04F27DBE870FBD7FE9D" box="[937,1038,316,337]" italics="true" pageId="16" pageNumber="1204">Discopyge</emphasis>
</taxonomicName>
</th>
<th id="76A05DBE22654FA02009E877FB17FE9D" box="[1147,1230,315,337]" gridcol="4" gridrow="0" pageId="16" pageNumber="1204">
<taxonomicName id="4C414D032265B04F200CE870FB12FE9D" baseAuthorityName="Claeson" baseAuthorityYear="2014" box="[1150,1227,316,337]" class="Elasmobranchii" family="Narcinidae" genus="Narcine" kingdom="Animalia" order="Torpediniformes" pageId="16" pageNumber="1204" phylum="Chordata" rank="genus">
<emphasis id="B935EA922265B04F200CE870FB12FE9D" box="[1150,1227,316,337]" italics="true" pageId="16" pageNumber="1204">Narcine</emphasis>
</taxonomicName>
</th>
<th id="76A05DBE22654FA02149E877FA63FE9D" box="[1339,1466,315,337]" gridcol="5" gridrow="0" pageId="16" pageNumber="1204">
<taxonomicName id="4C414D032265B04F2137E870FA63FE9D" authority="Trunk" authorityName="Trunk" box="[1349,1466,316,337]" class="Elasmobranchii" family="Narcinidae" genus="Titanonarke" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="16" pageNumber="1204" phylum="Chordata" rank="genus">
<emphasis id="B935EA922265B04F2137E870FA63FE9D" box="[1349,1466,316,337]" italics="true" pageId="16" pageNumber="1204">Titanonarke</emphasis>
</taxonomicName>
</th>
</tr>
<tr id="357134C222654FA024EDE821FA63FE4E" box="[159,1466,365,386]" gridrow="1" pageId="16" pageNumber="1204">
<th id="76A05DBE22654FA024EDE821FE52FE4E" box="[159,395,365,386]" gridcol="0" gridrow="1" pageId="16" pageNumber="1204">Trunk vertebrae</th>
<td id="76A05DBE22654FA0258AE821FDB2FE4E" box="[504,619,365,386]" gridcol="1" gridrow="1" pageId="16" pageNumber="1204">1320</td>
<td id="76A05DBE22654FA026A5E821FCE5FE4E" box="[727,828,365,386]" gridcol="2" gridrow="1" pageId="16" pageNumber="1204">2325</td>
<td id="76A05DBE22654FA027DBE821FBD7FE4E" box="[937,1038,365,386]" gridcol="3" gridrow="1" pageId="16" pageNumber="1204">20</td>
<td id="76A05DBE22654FA02009E821FB17FE4E" box="[1147,1230,365,386]" gridcol="4" gridrow="1" pageId="16" pageNumber="1204">1531</td>
<td id="76A05DBE22654FA02149E821FA63FE4E" box="[1339,1466,365,386]" gridcol="5" gridrow="1" pageId="16" pageNumber="1204">2730</td>
</tr>
<tr id="357134C222654FA024EDE8C1FA63FE6E" box="[159,1466,397,418]" gridrow="2" pageId="16" pageNumber="1204">
<th id="76A05DBE22654FA024EDE8C1FE52FE6E" box="[159,395,397,418]" gridcol="0" gridrow="2" pageId="16" pageNumber="1204">Precaudal vertebrae</th>
<td id="76A05DBE22654FA0258AE8C1FDB2FE6E" box="[504,619,397,418]" gridcol="1" gridrow="2" pageId="16" pageNumber="1204">4673</td>
<td id="76A05DBE22654FA026A5E8C1FCE5FE6E" box="[727,828,397,418]" gridcol="2" gridrow="2" pageId="16" pageNumber="1204">6163</td>
<td id="76A05DBE22654FA027DBE8C1FBD7FE6E" box="[937,1038,397,418]" gridcol="3" gridrow="2" pageId="16" pageNumber="1204">5870</td>
<td id="76A05DBE22654FA02009E8C1FB17FE6E" box="[1147,1230,397,418]" gridcol="4" gridrow="2" pageId="16" pageNumber="1204">5877</td>
<td id="76A05DBE22654FA02149E8C1FA63FE6E" box="[1339,1466,397,418]" gridcol="5" gridrow="2" pageId="16" pageNumber="1204">74115</td>
</tr>
<tr id="357134C222654FA024EDE8E1FA63FE0E" box="[159,1466,429,450]" gridrow="3" pageId="16" pageNumber="1204">
<th id="76A05DBE22654FA024EDE8E1FE52FE0E" box="[159,395,429,450]" gridcol="0" gridrow="3" pageId="16" pageNumber="1204">Caudal vertebrae</th>
<td id="76A05DBE22654FA0258AE8E1FDB2FE0E" box="[504,619,429,450]" gridcol="1" gridrow="3" pageId="16" pageNumber="1204">3146</td>
<td id="76A05DBE22654FA026A5E8E1FCE5FE0E" box="[727,828,429,450]" gridcol="2" gridrow="3" pageId="16" pageNumber="1204">2224</td>
<td id="76A05DBE22654FA027DBE8E1FBD7FE0E" box="[937,1038,429,450]" gridcol="3" gridrow="3" pageId="16" pageNumber="1204">?</td>
<td id="76A05DBE22654FA02009E8E1FB17FE0E" box="[1147,1230,429,450]" gridcol="4" gridrow="3" pageId="16" pageNumber="1204">1832</td>
<td id="76A05DBE22654FA02149E8E1FA63FE0E" box="[1339,1466,429,450]" gridcol="5" gridrow="3" pageId="16" pageNumber="1204">2332</td>
</tr>
<tr id="357134C222654FA024EDE881FA63FE2E" box="[159,1466,461,482]" gridrow="4" pageId="16" pageNumber="1204">
<th id="76A05DBE22654FA024EDE881FE52FE2E" box="[159,395,461,482]" gridcol="0" gridrow="4" pageId="16" pageNumber="1204">Total vertebrae</th>
<td id="76A05DBE22654FA0258AE881FDB2FE2E" box="[504,619,461,482]" gridcol="1" gridrow="4" pageId="16" pageNumber="1204">96118</td>
<td id="76A05DBE22654FA026A5E881FCE5FE2E" box="[727,828,461,482]" gridcol="2" gridrow="4" pageId="16" pageNumber="1204">97112</td>
<td id="76A05DBE22654FA027DBE881FBD7FE2E" box="[937,1038,461,482]" gridcol="3" gridrow="4" pageId="16" pageNumber="1204">8591</td>
<td id="76A05DBE22654FA02009E881FB17FE2E" box="[1147,1230,461,482]" gridcol="4" gridrow="4" pageId="16" pageNumber="1204">100127</td>
<td id="76A05DBE22654FA02149E881FA63FE2E" box="[1339,1466,461,482]" gridcol="5" gridrow="4" pageId="16" pageNumber="1204">133155</td>
</tr>
<tr id="357134C222654FA024EDE8A1FA63FDCE" box="[159,1466,493,514]" gridrow="5" pageId="16" pageNumber="1204">
<th id="76A05DBE22654FA024EDE8A1FE52FDCE" box="[159,395,493,514]" gridcol="0" gridrow="5" pageId="16" pageNumber="1204">Rib pairs</th>
<td id="76A05DBE22654FA0258AE8A1FDB2FDCE" box="[504,619,493,514]" gridcol="1" gridrow="5" pageId="16" pageNumber="1204">4</td>
<td id="76A05DBE22654FA026A5E8A1FCE5FDCE" box="[727,828,493,514]" gridcol="2" gridrow="5" pageId="16" pageNumber="1204">79</td>
<td id="76A05DBE22654FA027DBE8A1FBD7FDCE" box="[937,1038,493,514]" gridcol="3" gridrow="5" pageId="16" pageNumber="1204">8</td>
<td id="76A05DBE22654FA02009E8A1FB17FDCE" box="[1147,1230,493,514]" gridcol="4" gridrow="5" pageId="16" pageNumber="1204">510</td>
<td id="76A05DBE22654FA02149E8A1FA63FDCE" box="[1339,1466,493,514]" gridcol="5" gridrow="5" pageId="16" pageNumber="1204">810</td>
</tr>
<tr id="357134C222654FA024EDEB40FA63FDED" box="[159,1466,524,545]" gridrow="6" pageId="16" pageNumber="1204">
<th id="76A05DBE22654FA024EDEB40FE52FDED" box="[159,395,524,545]" gridcol="0" gridrow="6" pageId="16" pageNumber="1204">Total pectoral radials</th>
<td id="76A05DBE22654FA0258AEB40FDB2FDED" box="[504,619,524,545]" gridcol="1" gridrow="6" pageId="16" pageNumber="1204">?</td>
<td id="76A05DBE22654FA026A5EB40FCE5FDED" box="[727,828,524,545]" gridcol="2" gridrow="6" pageId="16" pageNumber="1204">31</td>
<td id="76A05DBE22654FA027DBEB40FBD7FDED" box="[937,1038,524,545]" gridcol="3" gridrow="6" pageId="16" pageNumber="1204">?</td>
<td id="76A05DBE22654FA02009EB40FB17FDED" box="[1147,1230,524,545]" gridcol="4" gridrow="6" pageId="16" pageNumber="1204">2741</td>
<td id="76A05DBE22654FA02149EB40FA63FDED" box="[1339,1466,524,545]" gridcol="5" gridrow="6" pageId="16" pageNumber="1204">3540</td>
</tr>
<tr id="357134C222654FA024EDEB60FA63FD8D" box="[159,1466,556,577]" gridrow="7" pageId="16" pageNumber="1204">
<th id="76A05DBE22654FA024EDEB60FE52FD8D" box="[159,395,556,577]" gridcol="0" gridrow="7" pageId="16" pageNumber="1204">Pelvic fin radials</th>
<td id="76A05DBE22654FA0258AEB60FDB2FD8D" box="[504,619,556,577]" gridcol="1" gridrow="7" pageId="16" pageNumber="1204">1213</td>
<td id="76A05DBE22654FA026A5EB60FCE5FD8D" box="[727,828,556,577]" gridcol="2" gridrow="7" pageId="16" pageNumber="1204">1621</td>
<td id="76A05DBE22654FA027DBEB60FBD7FD8D" box="[937,1038,556,577]" gridcol="3" gridrow="7" pageId="16" pageNumber="1204">?</td>
<td id="76A05DBE22654FA02009EB60FB17FD8D" box="[1147,1230,556,577]" gridcol="4" gridrow="7" pageId="16" pageNumber="1204">1421</td>
<td id="76A05DBE22654FA02149EB60FA63FD8D" box="[1339,1466,556,577]" gridcol="5" gridrow="7" pageId="16" pageNumber="1204">1921</td>
</tr>
<tr id="357134C222654FA024EDEB00FA63FDAD" box="[159,1466,588,609]" gridrow="8" pageId="16" pageNumber="1204">
<th id="76A05DBE22654FA024EDEB00FE52FDAD" box="[159,395,588,609]" gridcol="0" gridrow="8" pageId="16" pageNumber="1204">First dorsal fin radials</th>
<td id="76A05DBE22654FA0258AEB00FDB2FDAD" box="[504,619,588,609]" gridcol="1" gridrow="8" pageId="16" pageNumber="1204">56</td>
<td id="76A05DBE22654FA026A5EB00FCE5FDAD" box="[727,828,588,609]" gridcol="2" gridrow="8" pageId="16" pageNumber="1204">67</td>
<td id="76A05DBE22654FA027DBEB00FBD7FDAD" box="[937,1038,588,609]" gridcol="3" gridrow="8" pageId="16" pageNumber="1204">?</td>
<td id="76A05DBE22654FA02009EB00FB17FDAD" box="[1147,1230,588,609]" gridcol="4" gridrow="8" pageId="16" pageNumber="1204">610</td>
<td id="76A05DBE22654FA02149EB00FA63FDAD" box="[1339,1466,588,609]" gridcol="5" gridrow="8" pageId="16" pageNumber="1204">79</td>
</tr>
<tr id="357134C222654FA024EDEB20FA63FD4D" box="[159,1466,620,641]" gridrow="9" pageId="16" pageNumber="1204">
<th id="76A05DBE22654FA024EDEB20FE52FD4D" box="[159,395,620,641]" gridcol="0" gridrow="9" pageId="16" pageNumber="1204">Second dorsal fin radials</th>
<td id="76A05DBE22654FA0258AEB20FDB2FD4D" box="[504,619,620,641]" gridcol="1" gridrow="9" pageId="16" pageNumber="1204">67</td>
<td id="76A05DBE22654FA026A5EB20FCE5FD4D" box="[727,828,620,641]" gridcol="2" gridrow="9" pageId="16" pageNumber="1204">78</td>
<td id="76A05DBE22654FA027DBEB20FBD7FD4D" box="[937,1038,620,641]" gridcol="3" gridrow="9" pageId="16" pageNumber="1204">?</td>
<td id="76A05DBE22654FA02009EB20FB17FD4D" box="[1147,1230,620,641]" gridcol="4" gridrow="9" pageId="16" pageNumber="1204">611</td>
<td id="76A05DBE22654FA02149EB20FA63FD4D" box="[1339,1466,620,641]" gridcol="5" gridrow="9" pageId="16" pageNumber="1204">67</td>
</tr>
<tr id="357134C222654FA024EDEBC0FA63FD6D" box="[159,1466,652,673]" gridrow="10" pageId="16" pageNumber="1204">
<th id="76A05DBE22654FA024EDEBC0FE52FD6D" box="[159,395,652,673]" gridcol="0" gridrow="10" pageId="16" pageNumber="1204">Total caudal fin radials</th>
<td id="76A05DBE22654FA0258AEBC0FDB2FD6D" box="[504,619,652,673]" gridcol="1" gridrow="10" pageId="16" pageNumber="1204">3435</td>
<td id="76A05DBE22654FA026A5EBC0FCE5FD6D" box="[727,828,652,673]" gridcol="2" gridrow="10" pageId="16" pageNumber="1204">3336</td>
<td id="76A05DBE22654FA027DBEBC0FBD7FD6D" box="[937,1038,652,673]" gridcol="3" gridrow="10" pageId="16" pageNumber="1204">?</td>
<td id="76A05DBE22654FA02009EBC0FB17FD6D" box="[1147,1230,652,673]" gridcol="4" gridrow="10" pageId="16" pageNumber="1204">3963</td>
<td id="76A05DBE22654FA02149EBC0FA63FD6D" box="[1339,1466,652,673]" gridcol="5" gridrow="10" pageId="16" pageNumber="1204">4142</td>
</tr>
<tr id="357134C222654FA024EDEBE0FA63FD0D" box="[159,1466,684,705]" gridrow="11" pageId="16" pageNumber="1204">
<th id="76A05DBE22654FA024EDEBE0FE52FD0D" box="[159,395,684,705]" gridcol="0" gridrow="11" pageId="16" pageNumber="1204">Upper jaw tooth rows</th>
<td id="76A05DBE22654FA0258AEBE0FDB2FD0D" box="[504,619,684,705]" gridcol="1" gridrow="11" pageId="16" pageNumber="1204">920</td>
<td id="76A05DBE22654FA026A5EBE0FCE5FD0D" box="[727,828,684,705]" gridcol="2" gridrow="11" pageId="16" pageNumber="1204">1422</td>
<td id="76A05DBE22654FA027DBEBE0FBD7FD0D" box="[937,1038,684,705]" gridcol="3" gridrow="11" pageId="16" pageNumber="1204">1020</td>
<td id="76A05DBE22654FA02009EBE0FB17FD0D" box="[1147,1230,684,705]" gridcol="4" gridrow="11" pageId="16" pageNumber="1204">1127</td>
<td id="76A05DBE22654FA02149EBE0FA63FD0D" box="[1339,1466,684,705]" gridcol="5" gridrow="11" pageId="16" pageNumber="1204">1213</td>
</tr>
<tr id="357134C222654FA024EDEB80FA63FD2D" box="[159,1466,716,737]" gridrow="12" pageId="16" pageNumber="1204">
<th id="76A05DBE22654FA024EDEB80FE52FD2D" box="[159,395,716,737]" gridcol="0" gridrow="12" pageId="16" pageNumber="1204">Lower jaw tooth rows</th>
<td id="76A05DBE22654FA0258AEB80FDB2FD2D" box="[504,619,716,737]" gridcol="1" gridrow="12" pageId="16" pageNumber="1204">922</td>
<td id="76A05DBE22654FA026A5EB80FCE5FD2D" box="[727,828,716,737]" gridcol="2" gridrow="12" pageId="16" pageNumber="1204">1422</td>
<td id="76A05DBE22654FA027DBEB80FBD7FD2D" box="[937,1038,716,737]" gridcol="3" gridrow="12" pageId="16" pageNumber="1204">1020</td>
<td id="76A05DBE22654FA02009EB80FB17FD2D" box="[1147,1230,716,737]" gridcol="4" gridrow="12" pageId="16" pageNumber="1204">830</td>
<td id="76A05DBE22654FA02149EB80FA63FD2D" box="[1339,1466,716,737]" gridcol="5" gridrow="12" pageId="16" pageNumber="1204">1517</td>
</tr>
</table>
</paragraph>
<paragraph id="8BFE36802265B04F24C8ED36FC72FB9D" blockId="16.[159,781,827,1934]" lastBlockId="16.[845,1467,827,1105]" pageId="16" pageNumber="1204">
The pelvic fins (
<figureCitation id="137A2A052265B04F2515ED36FE6AFB5D" box="[359,435,1146,1169]" captionStart="Figure 11" captionStartId="11.[159,227,701,722]" captionTargetBox="[365,1257,183,669]" captionTargetId="figure-384@11.[365,1261,183,669]" captionTargetPageId="11" captionText="Figure 11. Pelvic fins of †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site. A, MCSNV IG.VR.67290; B, reconstruction. Abbreviations: bas, basipterygia; ilp, iliac process; pel, pelvic processes; pub, puboischiadic bar; rad, pelvic radials. The arrowhead indicates the iliac process. Scale bars = 50 mm." figureDoi="http://doi.org/10.5281/zenodo.10883297" httpUri="https://zenodo.org/record/10883297/files/figure.png" pageId="16" pageNumber="1204">Fig. 11</figureCitation>
) are small and single-lobed, and their anterior margin is straight and barely overlapped by the posterior margin of the pectoral disc. The maximum width of the pelvic fins is about 50% of the pectoral disc width, whereas their anterior margin is about 24% of the disc length. The puboischiadic bar is robust and wide, with a slightly concave anterior margin. The presence of the puboischiadic foramina is difficult to detect. The prepelvic processes are long and straight, extending anteriorly almost to the level of the scapulocoracoid. They are wider distally than along the shaft, resembling the condition of all narcinids (Rincon
<emphasis id="B935EA922265B04F25A8EC94FDD3FA23" box="[474,522,1496,1519]" italics="true" pageId="16" pageNumber="1204">et al</emphasis>
. 2001, fig. 8;
<bibRefCitation id="EFD04B712265B04F26D6EC94FEAAF9C3" author="Fechhelm, J. D. &amp; McEachran, J. D." pageId="16" pageNumber="1204" pagination="171 - 209" refId="ref21620" refString="Fechhelm, J. D. &amp; McEachran, J. D. 1984. A revision of the electric ray genus Diplobatis with notes on the interrelationships of Narcinidae (Chondrichthyes, Torpediniformes). Bulletin of the Florida State Museum Biological Sciences, 29, 171 - 209." type="journal article" year="1984">Fechhelm &amp; McEachran 1984</bibRefCitation>
, figs 7, 16). The distal end of the prepelvic process was described as spatulate by
<bibRefCitation id="EFD04B712265B04F26D9EF54FF31F983" author="Carvalho, M. R. de" pageId="16" pageNumber="1204" refId="ref19921" refString="Carvalho, M. R. de. 2010. Morphology and phylogenetic relationships of the giant electric ray from the Eocene of Monte Bolca, Italy (Chondrichthyes: Torpediniformes). Pp. 183 - 198 in D. K. Elliot, J. G. Maisey, X. Yu &amp; D. Miao (eds) Morphology, phylogeny and paleobiogeography of fossil fishes. Verlag Dr. Friedrich Pfeil, Munich." type="book" year="2010">Carvalho (2010)</bibRefCitation>
. However, the margin of this structure, identified with an arrowhead on the holotypic specimen
<collectionCode id="ED50AE452265B04F26DBEF14FCD4F9A3" box="[681,781,1624,1647]" pageId="16" pageNumber="1204">MGP-PD</collectionCode>
26275/6 by
<bibRefCitation id="EFD04B712265B04F2553EF34FE10F943" author="Carvalho, M. R. de" box="[289,457,1656,1679]" pageId="16" pageNumber="1204" refId="ref19921" refString="Carvalho, M. R. de. 2010. Morphology and phylogenetic relationships of the giant electric ray from the Eocene of Monte Bolca, Italy (Chondrichthyes: Torpediniformes). Pp. 183 - 198 in D. K. Elliot, J. G. Maisey, X. Yu &amp; D. Miao (eds) Morphology, phylogeny and paleobiogeography of fossil fishes. Verlag Dr. Friedrich Pfeil, Munich." type="book" year="2010">Carvalho (2010</bibRefCitation>
, fig. 7), is not part of the prepelvic process. The iliac process, preserved only on the left side of
<collectionCode id="ED50AE452265B04F2555EFF4FE5CF903" box="[295,389,1720,1743]" pageId="16" pageNumber="1204">MCSNV</collectionCode>
<collectionCode id="ED50AE452265B04F25E3EFF4FE76F903" box="[401,431,1720,1743]" country="China" name="Institute of Geology" pageId="16" pageNumber="1204">IG</collectionCode>
.VR.67290, appears short, stout and straight (
<figureCitation id="137A2A052265B04F255DEF94FEA4F923" box="[303,381,1752,1775]" captionStart="Figure 11" captionStartId="11.[159,227,701,722]" captionTargetBox="[365,1257,183,669]" captionTargetId="figure-384@11.[365,1261,183,669]" captionTargetPageId="11" captionText="Figure 11. Pelvic fins of †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site. A, MCSNV IG.VR.67290; B, reconstruction. Abbreviations: bas, basipterygia; ilp, iliac process; pel, pelvic processes; pub, puboischiadic bar; rad, pelvic radials. The arrowhead indicates the iliac process. Scale bars = 50 mm." figureDoi="http://doi.org/10.5281/zenodo.10883297" httpUri="https://zenodo.org/record/10883297/files/figure.png" pageId="16" pageNumber="1204">Fig. 11</figureCitation>
) if compared to the long and curved iliac process of
<taxonomicName id="4C414D032265B04F252FEFBBFE6AF8C2" baseAuthorityName="Claeson" baseAuthorityYear="2014" box="[349,435,1783,1806]" class="Elasmobranchii" family="Narcinidae" genus="Narcine" kingdom="Animalia" order="Torpediniformes" pageId="16" pageNumber="1204" phylum="Chordata" rank="genus">
<emphasis id="B935EA922265B04F252FEFBBFE6AF8C2" box="[349,435,1783,1806]" italics="true" pageId="16" pageNumber="1204">Narcine</emphasis>
</taxonomicName>
and
<taxonomicName id="4C414D032265B04F2584EFBBFDBFF8C2" authorityName="Heckel" authorityYear="1846" box="[502,614,1783,1806]" class="Elasmobranchii" family="Narcinidae" genus="Discopyge" kingdom="Animalia" order="Torpediniformes" pageId="16" pageNumber="1204" phylum="Chordata" rank="genus">
<emphasis id="B935EA922265B04F2584EFBBFDBFF8C2" box="[502,614,1783,1806]" italics="true" pageId="16" pageNumber="1204">Discopyge</emphasis>
</taxonomicName>
(Menni
<emphasis id="B935EA922265B04F26A2EFBBFCDEF8C2" box="[720,775,1783,1806]" italics="true" pageId="16" pageNumber="1204">et al</emphasis>
. 2008;
<bibRefCitation id="EFD04B712265B04F2496EE5BFEAFF8E2" author="Claeson, K. M." box="[228,374,1815,1838]" pageId="16" pageNumber="1204" pagination="597 - 612" refId="ref20593" refString="Claeson, K. M. 2014. The impacts of comparative anatomy of electric rays (Batoidea: Torpediniformes) on their systematic hypotheses. Journal of Morphology, 275, 597 - 612." type="journal article" year="2014">Claeson 2014</bibRefCitation>
). The basipterygia are slightly longer than the puboischiadic bar, and have a slightly concave inner margin. Each basipterygium supports about 2124 pelvic fin radials, the first of which is enlarged, articulates with the lateral node of the puboischiadic bar and supports the anterior margin of the pelvic fin. Each pelvic fin radial bifurcates distally once and is composed of three segments. A single individual (
<collectionCode id="ED50AE452265B04F20F0EAD7FB3BFC7E" box="[1154,1250,923,946]" pageId="16" pageNumber="1204">MCSNV</collectionCode>
<collectionCode id="ED50AE452265B04F209EEAD7FAD3FC7E" box="[1260,1290,923,946]" country="China" name="Institute of Geology" pageId="16" pageNumber="1204">IG</collectionCode>
.91128/9) seems to show an elongate clasper, articulating with the distal tip of the basipterygium (
<figureCitation id="137A2A052265B04F2023EA96FB7CFC3D" box="[1105,1189,986,1009]" captionStart="Figure 12" captionStartId="11.[159,227,1780,1801]" captionTargetBox="[166,774,1083,1748]" captionTargetId="figure-435@11.[166,774,1083,1748]" captionTargetPageId="11" captionText="Figure 12. †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site. Close-up of the distal end of pelvic fins of MCSNV IG.91128, which is supposed to be the unique male individual based on the presence of claspers. Abbreviations: cla, clasper; dfr, first dorsal-fin radials, pelvic radials. Scale bar = 50 mm." figureDoi="http://doi.org/10.5281/zenodo.10883299" httpUri="https://zenodo.org/record/10883299/files/figure.png" pageId="16" pageNumber="1204">Fig. 12</figureCitation>
). In length, the clasper appears to extend past the posterior tip of the pelvic fin lobe. However, it was not possible to analyse this structure in detail.
</paragraph>
<paragraph id="8BFE36802265B04F2707ED26FAA3F9F3" blockId="16.[845,1467,1130,1599]" pageId="16" pageNumber="1204">
<emphasis id="B935EA922265B04F2707ED26FBDFFB4D" bold="true" box="[885,1030,1130,1153]" pageId="16" pageNumber="1204">Median fins.</emphasis>
There are two dorsal fins (
<figureCitation id="137A2A052265B04F2135ED26FA76FB4D" box="[1351,1455,1130,1153]" captionStart="Figure 13" captionStartId="12.[159,227,761,782]" captionTargetBox="[317,1309,183,729]" captionTargetId="figure-291@12.[317,1309,183,729]" captionTargetPageId="12" captionText="Figure 13. †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site. Details of the precaudal tail of A, MCSNV IG. VR.67290 and B, MGP-PD 26276. The arrows mark the position of the two dorsal fins in MCSNV IG.VR.67290 and the second dorsal fin in MGP-PD 26276 already detected and figured by Jaekel (1894). C, Detail of the caudal fin of MCSNV IG.91129. Scale bars: A, B = 50 mm; C = 10 mm." figureDoi="http://doi.org/10.5281/zenodo.10883301" httpUri="https://zenodo.org/record/10883301/files/figure.png" pageId="16" pageNumber="1204">Fig. 13A</figureCitation>
). The first one originates at about 62% SL, is slightly larger than the second one and is supported by seven to nine radials. The second dorsal fin originates at about 78% SL and is supported by six or seven radials. The interdorsal distance measures about 7% SL. The obvious presence of two dorsal fins in
<collectionCode id="ED50AE452265B04F205AEC65FB5FFA8C" box="[1064,1158,1321,1344]" pageId="16" pageNumber="1204">MCSNV</collectionCode>
<collectionCode id="ED50AE452265B04F20E1EC65FB69FA8C" box="[1171,1200,1321,1344]" country="China" name="Institute of Geology" pageId="16" pageNumber="1204">IG</collectionCode>
.VR.67290 supports the interpretation of
<bibRefCitation id="EFD04B712265B04F206BEC05FB6FFAAC" author="Jaekel, O." box="[1049,1206,1353,1376]" pageId="16" pageNumber="1204" refId="ref23357" refString="Jaekel, O. 1894. Die eoca ¨ nen Selachier vom Monte Bolca: ein Beitrag zur Morphogenie der Wirbelthiere. Springer, Berlin, 177 pp." type="book" year="1894">Jaekel (1894)</bibRefCitation>
and
<bibRefCitation id="EFD04B712265B04F2170EC05FA62FAAC" author="Cappetta, H." box="[1282,1467,1353,1376]" pageId="16" pageNumber="1204" refId="ref19193" refString="Cappetta, H. 2012. Handbook of Paleoichthyology - Chondrichthyes - Mesozoic and Cenozoic Elasmobranchii: Teeth. Verlag Dr. Friedrich Pfeil, Munich, 512 pp." type="book" year="2012">Cappetta (2012)</bibRefCitation>
regarding the presence of at least one dorsal fin in the
<typeStatus id="54FA88222265B04F273FECC5FC73FA6C" box="[845,938,1417,1440]" pageId="16" pageNumber="1204" type="holotype">holotype</typeStatus>
<collectionCode id="ED50AE452265B04F27C4ECC5FBC3FA6C" box="[950,1050,1417,1440]" pageId="16" pageNumber="1204">MGP-PD</collectionCode>
26275/6 (
<figureCitation id="137A2A052265B04F20E1ECC5FB21FA6C" box="[1171,1272,1417,1440]" captionStart="Figure 13" captionStartId="12.[159,227,761,782]" captionTargetBox="[317,1309,183,729]" captionTargetId="figure-291@12.[317,1309,183,729]" captionTargetPageId="12" captionText="Figure 13. †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site. Details of the precaudal tail of A, MCSNV IG. VR.67290 and B, MGP-PD 26276. The arrows mark the position of the two dorsal fins in MCSNV IG.VR.67290 and the second dorsal fin in MGP-PD 26276 already detected and figured by Jaekel (1894). C, Detail of the caudal fin of MCSNV IG.91129. Scale bars: A, B = 50 mm; C = 10 mm." figureDoi="http://doi.org/10.5281/zenodo.10883301" httpUri="https://zenodo.org/record/10883301/files/figure.png" pageId="16" pageNumber="1204">Fig. 13B</figureCitation>
). The inadequate preservation of the dorsal fins in the historical material prevented their recognition by
<bibRefCitation id="EFD04B712265B04F20CFEC84FAAEFA13" author="Carvalho, M. R. de" box="[1213,1399,1480,1503]" pageId="16" pageNumber="1204" refId="ref19921" refString="Carvalho, M. R. de. 2010. Morphology and phylogenetic relationships of the giant electric ray from the Eocene of Monte Bolca, Italy (Chondrichthyes: Torpediniformes). Pp. 183 - 198 in D. K. Elliot, J. G. Maisey, X. Yu &amp; D. Miao (eds) Morphology, phylogeny and paleobiogeography of fossil fishes. Verlag Dr. Friedrich Pfeil, Munich." type="book" year="2010">Carvalho (2010)</bibRefCitation>
, who erroneously regarded their absence as diagnostic for †
<taxonomicName id="4C414D032265B04F272AEF44FC03F9D3" box="[856,986,1544,1567]" class="Elasmobranchii" family="Narcinidae" genus="Titanonarke" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="16" pageNumber="1204" phylum="Chordata" rank="genus">
<emphasis id="B935EA922265B04F272AEF44FC03F9D3" box="[856,986,1544,1567]" italics="true" pageId="16" pageNumber="1204">Titanonarke</emphasis>
</taxonomicName>
. About 42 radials support the caudal fin, of which about 20 are dorsal and 22 ventral (
<figureCitation id="137A2A052265B04F217FEF64FAB4F9F3" box="[1293,1389,1576,1599]" captionStart="Figure 13" captionStartId="12.[159,227,761,782]" captionTargetBox="[317,1309,183,729]" captionTargetId="figure-291@12.[317,1309,183,729]" captionTargetPageId="12" captionText="Figure 13. †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site. Details of the precaudal tail of A, MCSNV IG. VR.67290 and B, MGP-PD 26276. The arrows mark the position of the two dorsal fins in MCSNV IG.VR.67290 and the second dorsal fin in MGP-PD 26276 already detected and figured by Jaekel (1894). C, Detail of the caudal fin of MCSNV IG.91129. Scale bars: A, B = 50 mm; C = 10 mm." figureDoi="http://doi.org/10.5281/zenodo.10883301" httpUri="https://zenodo.org/record/10883301/files/figure.png" pageId="16" pageNumber="1204">Fig. 13C</figureCitation>
).
</paragraph>
<paragraph id="8BFE36802265B04E2707EF14FE6EFE98" blockId="16.[845,1467,1624,1934]" lastBlockId="17.[159,781,189,659]" lastPageId="17" lastPageNumber="1205" pageId="16" pageNumber="1204">
<emphasis id="B935EA922265B04F2707EF14FC3FF9A3" bold="true" box="[885,998,1624,1647]" pageId="16" pageNumber="1204">Dentition.</emphasis>
The teeth of †
<taxonomicName id="4C414D032265B04F20FFEF14FAD6F9A3" box="[1165,1295,1624,1647]" class="Elasmobranchii" family="Narcinidae" genus="Titanonarke" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="16" pageNumber="1204" phylum="Chordata" rank="genus">
<emphasis id="B935EA922265B04F20FFEF14FAD6F9A3" box="[1165,1295,1624,1647]" italics="true" pageId="16" pageNumber="1204">Titanonarke</emphasis>
</taxonomicName>
are arranged in tooth bands medially across the jaw symphyses, forming a tessellated pavement (
<figureCitation id="137A2A052265B04F2031EFD4FB4AF963" box="[1091,1171,1688,1711]" captionStart="Figure 14" captionStartId="12.[159,227,1806,1827]" captionTargetBox="[166,774,1108,1775]" captionTargetId="figure-371@12.[166,774,1108,1775]" captionTargetPageId="12" captionText="Figure 14. †Titanonarke molini (Jaekel, 1894) from the Eocene of Monte Postale site. A, upper and lower tooth bands in MCSNV IG.VR.67290, with a close-up of some teeth in the area indicated. B, reconstruction. Abbreviations: la, labial cartilage; me, Meckels cartilage; pq, palatoquadrate. Scale bars 5 mm." figureDoi="http://doi.org/10.5281/zenodo.10883305" httpUri="https://zenodo.org/record/10883305/files/figure.png" pageId="16" pageNumber="1204">Fig. 14</figureCitation>
). The lower tooth band is wider than the upper one. It is not possible to detect the tooth formula, but the teeth of †
<taxonomicName id="4C414D032265B04F20CFEF94FAFBF923" baseAuthorityName="Jaekel" baseAuthorityYear="1894" box="[1213,1314,1752,1775]" class="Elasmobranchii" family="Narcinidae" genus="Titanonarke" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="16" pageNumber="1204" phylum="Chordata" rank="species" species="molini">
<emphasis id="B935EA922265B04F20CFEF94FAFBF923" box="[1213,1314,1752,1775]" italics="true" pageId="16" pageNumber="1204">T. molini</emphasis>
</taxonomicName>
appear to be arranged in at least 1213 rows in the upper jaw and 15 16 rows in the lower jaw, counted on symphyseal tooth series. The dentition is gradient monognatic heterodont with lateral and posterior tooth crowns becoming slightly lower. However, both upper and lower jaws show very little heterodonty. Sexual heterodonty (studied via the analysis of the specimens with and without claspers) appears to be absent. It is not possible to detect any ontogenetic heterodonty due to the poor-quality preservation of this region in juveniles.
</paragraph>
<paragraph id="8BFE36802264B04E24C8E811FED8FD5F" blockId="17.[159,781,189,659]" pageId="17" pageNumber="1205">
The tooth morphology is generally consistent with that of
<taxonomicName id="4C414D032264B04E24CCE831FECDFE58" baseAuthorityName="Claeson" baseAuthorityYear="2014" box="[190,276,381,404]" class="Elasmobranchii" family="Narcinidae" genus="Narcine" kingdom="Animalia" order="Torpediniformes" pageId="17" pageNumber="1205" phylum="Chordata" rank="genus">
<emphasis id="B935EA922264B04E24CCE831FECDFE58" box="[190,276,381,404]" italics="true" pageId="17" pageNumber="1205">Narcine</emphasis>
</taxonomicName>
(see
<bibRefCitation id="EFD04B712264B04E253DE831FDF8FE58" author="Herman, J. &amp; Hovestadt-Euler, M. &amp; Hovestadt, D. C. &amp; Stehmann, M." box="[335,545,381,404]" pageId="17" pageNumber="1205" pagination="5 - 45" refId="ref22824" refString="Herman, J., Hovestadt-Euler, M., Hovestadt, D. C. &amp; Stehmann, M. 2002. Contributions to the study of the comparative morphology of teeth and other relevant ichthyodorulites in living supraspecific taxa of chondrichthyan fishes. Part B: Batomorphii. No. 4: Order Torpediniformes - Family Narcinidae - Subfamily Narcininae - Genera: Benthobatis, Diplobatis, Discopyge and Narcine, Subfamily Narkinae - Genera: Bengalichthys, Crassinarke, Heteronarce, Narke, Temera, and Typhlonarke, Family Torpedinidae - Subfamily Torpedininae - Genus: Torpedo - Subgenus: T. (Tetronarke) and T. (Torpedo) and Subfamily Hypninae - Genus: Hypnos. Bulletin de l'Institut Royal des Sciences Naturelles de Belgique, Biologie, 72, 5 - 45." type="journal article" year="2002">
Herman
<emphasis id="B935EA922264B04E25DFE831FE04FE58" box="[429,477,381,404]" italics="true" pageId="17" pageNumber="1205">et al</emphasis>
. 2002
</bibRefCitation>
, pls 10, 11). A single narrow, high and subtriangular cusp is present in each tooth. There are no accessory lateral cusplets. The crown base is broad and subcircular, and wider than the cusp length. The width of the cusp is less than half the length of the cutting edges. Cutting edges are blade-like. Lingual and labial ornamentations are absent and the tooth crown is completely smooth. Some teeth display a high and narrow root.
</paragraph>
<paragraph id="8BFE36802264B04E24B5EBE0FE42F9F3" blockId="17.[159,782,684,1599]" pageId="17" pageNumber="1205">
<emphasis id="B935EA922264B04E24B5EBE0FEBBFD0F" bold="true" box="[199,354,684,707]" pageId="17" pageNumber="1205">Gut contents.</emphasis>
The
<typeStatus id="54FA88222264B04E25D1EBE0FDD9FD0F" box="[419,512,684,707]" pageId="17" pageNumber="1205" type="holotype">holotype</typeStatus>
of †
<taxonomicName id="4C414D032264B04E2647EBE0FD4EFD0F" baseAuthorityName="Jaekel" baseAuthorityYear="1894" box="[565,663,684,707]" class="Elasmobranchii" family="Narcinidae" genus="Titanonarke" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="17" pageNumber="1205" phylum="Chordata" rank="species" species="molini">
<emphasis id="B935EA922264B04E2647EBE0FD4EFD0F" box="[565,663,684,707]" italics="true" pageId="17" pageNumber="1205">T. molini</emphasis>
</taxonomicName>
(
<collectionCode id="ED50AE452264B04E26D5EBE0FCD4FD0F" box="[679,781,684,707]" pageId="17" pageNumber="1205">MGP-PD</collectionCode>
26275/6) shows abdominal contents consisting of a pelletlike accumulation of hundreds of specimens of †
<taxonomicName id="4C414D032264B04E26D5EBA7FCD4FCCE" authorityName="d'Orbigny" authorityYear="1826" box="[679,781,747,770]" class="Magnoliopsida" family="Loranthaceae" genus="Alveolina" kingdom="Plantae" order="Santalales" pageId="17" pageNumber="1205" phylum="Tracheophyta" rank="genus">
<emphasis id="B935EA922264B04E26D5EBA7FCD4FCCE" box="[679,781,747,770]" italics="true" pageId="17" pageNumber="1205">Alveolina</emphasis>
</taxonomicName>
(
<figureCitation id="137A2A052264B04E24DAEA47FEF7FCEE" box="[168,302,779,802]" captionStart="Figure 15" captionStartId="13.[159,227,1162,1183]" captionTargetBox="[173,1451,183,1130]" captionTargetId="figure-322@13.[173,1453,183,1130]" captionTargetPageId="13" captionText="Figure 15. A, B, †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site; A, detail of the abdominal region in MGP-PD 26275 showing the stomach content; B, reconstruction; note also the embryo lying next to the stomach. C, D, close-up of some of the larger foraminifera of the genus †Alveolina in the stomach of MGP-PD 26275. E, dissected specimen of Torpedo nobiliana in ventral view (ESB tn200707_159) showing the position of the stomach, used to identify the accumulation in MGP-PD 26275 as gut contents. Abbreviations: int, intestine; liv, liver; met, metapterygium; pub, puboischiadic bar; sca, scapulocoracoid; st, stomach, syn synarcual. Scale bars: A, B = 50 mm; C, D = 2 mm." figureDoi="http://doi.org/10.5281/zenodo.10883307" httpUri="https://zenodo.org/record/10883307/files/figure.png" pageId="17" pageNumber="1205">Fig. 15AD</figureCitation>
), which is the most common foraminiferan genus in the Monte Postale sediments (Papazzoni
<emphasis id="B935EA922264B04E26A1EA67FCDEFC8E" box="[723,775,811,834]" italics="true" pageId="17" pageNumber="1205">et al</emphasis>
. 2017). The individual foraminifera are grouped together and closely packed, forming an accumulation, which is ovoid in outline and anteroposteriorly elongate, measuring about 80.5 and 41.8 mm in length and width, respectively. The exceptionally preserved gut contents show little evidence of digestion, suggesting that consumption occurred shortly before the numbfishs death. The accumulation is almost totally preserved in the abdominal cavity between pectoral and pelvic girdles, on one side of the vertebral axis, and just posteriorly to the flanges of the synarcual cartilage. This position is exactly comparable to that occupied by the gut-intestine tract in living electric rays (
<figureCitation id="137A2A052264B04E24DAED85FED2FB2C" box="[168,267,1225,1248]" captionStart="Figure 15" captionStartId="13.[159,227,1162,1183]" captionTargetBox="[173,1451,183,1130]" captionTargetId="figure-322@13.[173,1453,183,1130]" captionTargetPageId="13" captionText="Figure 15. A, B, †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site; A, detail of the abdominal region in MGP-PD 26275 showing the stomach content; B, reconstruction; note also the embryo lying next to the stomach. C, D, close-up of some of the larger foraminifera of the genus †Alveolina in the stomach of MGP-PD 26275. E, dissected specimen of Torpedo nobiliana in ventral view (ESB tn200707_159) showing the position of the stomach, used to identify the accumulation in MGP-PD 26275 as gut contents. Abbreviations: int, intestine; liv, liver; met, metapterygium; pub, puboischiadic bar; sca, scapulocoracoid; st, stomach, syn synarcual. Scale bars: A, B = 50 mm; C, D = 2 mm." figureDoi="http://doi.org/10.5281/zenodo.10883307" httpUri="https://zenodo.org/record/10883307/files/figure.png" pageId="17" pageNumber="1205">Fig. 15E</figureCitation>
), and in narcinids in particular (see
<bibRefCitation id="EFD04B712264B04E26DAED85FEC3FACC" author="Marinsek, G. P. &amp; Viliod, M. C. L. &amp; Mari, R. B." pageId="17" pageNumber="1205" pagination="229 - 236" refId="ref24058" refString="Marinsek, G. P., Viliod, M. C. L. &amp; Mari, R. B. 2017. Ecomorphology of the digestive tract of the brazilian electric ray Narcine brasiliensis (Olfers, 1831) (Torpediniformes: Narcinidae). Acta Zoologica, 98, 229 - 236." type="journal article" year="2017">
Marinsek
<emphasis id="B935EA922264B04E24EDEDA5FF0AFACC" box="[159,211,1257,1280]" italics="true" pageId="17" pageNumber="1205">et al</emphasis>
. 2017
</bibRefCitation>
, fig. 1). Moreover, the general shape of the accumulation resembles fossilized gastric contents detected in other elasmobranchs (e.g.
<bibRefCitation id="EFD04B712264B04E2641EC65FE95FAAC" author="Hovestadt, D. C. &amp; Hovestadt-Euler, M." pageId="17" pageNumber="1205" pagination="171 - 182" refId="ref23116" refString="Hovestadt, D. C. &amp; Hovestadt-Euler, M. 2002. The remains of a carcharhinid shark with a new triakid species in its digestive tract from the Oligocene of Germany. Tertiary Research, 21 (1 - 4), 171 - 182." type="journal article" year="2002">Hovestadt &amp; Hovestadt-Euler 2002</bibRefCitation>
;
<bibRefCitation id="EFD04B712264B04E2513EC05FDBAFAAC" author="Amalfitano, J. &amp; Dalla Vecchia, F. M. &amp; Giusberti, L. &amp; Fornaciari, E. &amp; Luciani, V. &amp; Roghi, G." box="[353,611,1353,1376]" pageId="17" pageNumber="1205" pagination="104 - 121" refId="ref17957" refString="Amalfitano, J., Dalla Vecchia, F. M., Giusberti, L., Fornaciari, E., Luciani, V. &amp; Roghi, G. 2017. Direct evidence of trophic interaction between a large lamniform shark, Cretodus sp., and a marine turtle from the Cretaceous of northeastern Italy. Palaeogeography, Palaeoclimatology, Palaeoecology, 469, 104 - 121." type="journal article" year="2017">
Amalfitano
<emphasis id="B935EA922264B04E2597EC05FDC2FAAC" box="[485,539,1353,1376]" italics="true" pageId="17" pageNumber="1205">et al</emphasis>
. 2017
</bibRefCitation>
). Additionally, there is no evidence of skeletal dispersal due to currents or bioturbation. Thus, we conclude that the accumulation is not the result of processes related to bottom tractive currents or bioturbation that sometimes can result in the accumulation of such elements (see Sch¨afer 1972). Consequently, this accumulation of foraminifera unquestionably represents gut contents.
</paragraph>
<paragraph id="8BFE36802264B04E24EDEF14FB44FD5F" blockId="17.[159,782,1624,1934]" lastBlockId="17.[845,1467,189,1488]" pageId="17" pageNumber="1205">
<emphasis id="B935EA922264B04E24EDEF14FEDAF9A3" bold="true" box="[159,259,1624,1647]" pageId="17" pageNumber="1205">Embryo.</emphasis>
MGP-UP 26275/6 also shows a unique partially developed embryo whose length is estimated to range between 50 and
<quantity id="4CB99B652264B04E253CEFD4FE40F963" box="[334,409,1688,1711]" metricMagnitude="-2" metricUnit="m" metricValue="6.0" pageId="17" pageNumber="1205" unit="mm" value="60.0">60 mm</quantity>
(
<figureCitation id="137A2A052264B04E25DBEFD4FE2CF963" box="[425,501,1688,1711]" captionStart="Figure 16" captionStartId="14.[159,227,1045,1066]" captionTargetBox="[173,1453,183,1013]" captionTargetId="figure-382@14.[173,1453,183,1013]" captionTargetPageId="14" captionText="Figure 16. A, B, D, †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site; A, close-up of the abdominal region of MGP-PD 26275 showing the embryo; the anterior region of the body lies on the lower portion of the photo; B, reconstruction; D, detail of the vertebral column of the embryo indicated in B with a dotted rectangle. C, dissected specimen of Potamotrygon tigrina (IUWP 7361) showing the position of the left uterus, just next to the stomach. Abbreviations: int, intestine; liv, liver; lu, left uterus; na, neural arches; ru, right uterus; st, stomach; vc, vertebral centra. Scale bars: A, B = 10 mm; D = 1 mm." figureDoi="http://doi.org/10.5281/zenodo.10883311" httpUri="https://zenodo.org/record/10883311/files/figure.png" pageId="17" pageNumber="1205">Fig. 16</figureCitation>
), about 67% of the adult size. The embryo comprises an almost complete vertebral column having about 150 vertebrae, whose number is perfectly comparable to that of an adult individual of †
<taxonomicName id="4C414D032264B04E268AEFBBFF3AF8E2" baseAuthorityName="Jaekel" baseAuthorityYear="1894" class="Elasmobranchii" family="Narcinidae" genus="Titanonarke" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="17" pageNumber="1205" phylum="Chordata" rank="species" species="molini">
<emphasis id="B935EA922264B04E268AEFBBFF3AF8E2" italics="true" pageId="17" pageNumber="1205">T. molini</emphasis>
</taxonomicName>
. The vertebral column is partially disarticulated, with some elements scattered from their original position. Each vertebra consists of a vertebral centrum and associated neural arch. There is no trace of the synarcual cartilage. This is consistent with the pattern of mineralization present in extant developing batoid embryos (e.g.
<taxonomicName id="4C414D032264B04E273FE9B1FC66FED8" box="[845,959,253,276]" class="Elasmobranchii" family="Myliobatidae" genus="Myliobatis" kingdom="Animalia" order="Myliobatiformes" pageId="17" pageNumber="1205" phylum="Chordata" rank="genus">
<emphasis id="B935EA922264B04E273FE9B1FC66FED8" box="[845,959,253,276]" italics="true" pageId="17" pageNumber="1205">Myliobatis</emphasis>
</taxonomicName>
,
<taxonomicName id="4C414D032264B04E27BFE9B1FBCBFED8" box="[973,1042,253,276]" class="Elasmobranchii" family="Pristidae" genus="Pristis" kingdom="Animalia" order="Rhinopristiformes" pageId="17" pageNumber="1205" phylum="Chordata" rank="genus">
<emphasis id="B935EA922264B04E27BFE9B1FBCBFED8" box="[973,1042,253,276]" italics="true" pageId="17" pageNumber="1205">Pristis</emphasis>
</taxonomicName>
,
<taxonomicName id="4C414D032264B04E2052E9B1FB8BFED8" authorityName="Linnaeus" authorityYear="1758" box="[1056,1106,253,276]" class="Elasmobranchii" family="Rajidae" genus="Raja" kingdom="Animalia" order="Rajiformes" pageId="17" pageNumber="1205" phylum="Chordata" rank="genus">
<emphasis id="B935EA922264B04E2052E9B1FB8BFED8" box="[1056,1106,253,276]" italics="true" pageId="17" pageNumber="1205">Raja</emphasis>
</taxonomicName>
; KMC pers. obs.), where the areolar cartilage of the free vertebrae is well mineralized earlier than the tessellated cartilage of the synarcual. The vertebral column is the only fossilized structure in this embryo. There are no cranial or girdle skeletal elements preserved. This condition is consistent with the late phases of skeletogenesis in elasmobranch fishes in which the mineralization of the cartilages involves only teeth, dermal denticles, vertebral centra and neural arches in very early stages (see e.g.
<bibRefCitation id="EFD04B712264B04E2035EB50FAC1FDFF" author="Eames, B. F. &amp; Allen, N. &amp; Young, J. &amp; Kaplan, A. &amp; Helms, J. A. &amp; Schneider, R. A." box="[1095,1304,540,563]" pageId="17" pageNumber="1205" pagination="542 - 554" refId="ref21336" refString="Eames, B. F., Allen, N., Young, J., Kaplan, A., Helms, J. A. &amp; Schneider, R. A. 2007. Skeletogenesis in the swell shark Cephaloscyllium ventriosum. Journal of Anatomy, 210, 542 - 554." type="journal article" year="2007">
Eames
<emphasis id="B935EA922264B04E20E9EB50FB09FDFF" box="[1179,1232,540,563]" italics="true" pageId="17" pageNumber="1205">et al</emphasis>
. 2007
</bibRefCitation>
;
<bibRefCitation id="EFD04B712264B04E215FEB50FC5BFD9F" author="Enault, S. &amp; Adnet, S. &amp; Debiais-Thibaud, M." pageId="17" pageNumber="1205" pagination="2" refId="ref21458" refString="Enault, S., Adnet, S. &amp; Debiais-Thibaud, M. 2016. Skeletogenesis during the late embryonic development of the catshark Scyliorhinus canicula (Chondrichthyes, Neoselachii). MorphoMuseuM, 1 (4), e 2, doi: 10.18563 / m 3.1.4. e 2" type="journal article" year="2016">
Enault
<emphasis id="B935EA922264B04E210DEB50FA6DFDFF" box="[1407,1460,540,563]" italics="true" pageId="17" pageNumber="1205">et al</emphasis>
. 2016
</bibRefCitation>
). However, teeth are not clearly recognizable in the embryo, whereas dermal denticles are expected to be absent, as in all torpediniforms.
</paragraph>
<paragraph id="8BFE36802264B04E2715EBD0FACBFA1C" blockId="17.[845,1467,189,1488]" pageId="17" pageNumber="1205">
The embryo is totally preserved in the abdominal cavity between the pectoral and pelvic girdle, on the left side of the vertebral axis, and just next to the stomach (see also
<figureCitation id="137A2A052264B04E273FEBB7FC43FCDE" box="[845,922,763,786]" captionStart="Figure 15" captionStartId="13.[159,227,1162,1183]" captionTargetBox="[173,1451,183,1130]" captionTargetId="figure-322@13.[173,1453,183,1130]" captionTargetPageId="13" captionText="Figure 15. A, B, †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site; A, detail of the abdominal region in MGP-PD 26275 showing the stomach content; B, reconstruction; note also the embryo lying next to the stomach. C, D, close-up of some of the larger foraminifera of the genus †Alveolina in the stomach of MGP-PD 26275. E, dissected specimen of Torpedo nobiliana in ventral view (ESB tn200707_159) showing the position of the stomach, used to identify the accumulation in MGP-PD 26275 as gut contents. Abbreviations: int, intestine; liv, liver; met, metapterygium; pub, puboischiadic bar; sca, scapulocoracoid; st, stomach, syn synarcual. Scale bars: A, B = 50 mm; C, D = 2 mm." figureDoi="http://doi.org/10.5281/zenodo.10883307" httpUri="https://zenodo.org/record/10883307/files/figure.png" pageId="17" pageNumber="1205">Fig. 15</figureCitation>
). This position is totally comparable to that occupied by the left uterus in fossil (
<bibRefCitation id="EFD04B712264B04E20E9EA57FAAAFCFE" author="Carvalho, M. R. de &amp; Maisey, J. C. &amp; Grande, L." box="[1179,1395,795,818]" pageId="17" pageNumber="1205" pagination="1 - 136" refId="ref20349" refString="Carvalho, M. R. de, Maisey, J. C. &amp; Grande, L. 2004. Freshwater stingrays of the Green River Formation of Wyoming (Early Eocene), with the description of a new genus and species and an analysis of its phylogenetic relationships (Chondrichthyes: Myliobatiformes). Bulletin of the American Museum of Natural History, 284, 1 - 136." type="journal article" year="2004">
Carvalho
<emphasis id="B935EA922264B04E2171EA57FAE8FCFE" box="[1283,1329,795,818]" italics="true" pageId="17" pageNumber="1205">et al</emphasis>
. 2004
</bibRefCitation>
, figs 2 and 13) and living batoids (
<figureCitation id="137A2A052264B04E2008EA77FB03FC9E" box="[1146,1242,827,850]" captionStart="Figure 16" captionStartId="14.[159,227,1045,1066]" captionTargetBox="[173,1453,183,1013]" captionTargetId="figure-382@14.[173,1453,183,1013]" captionTargetPageId="14" captionText="Figure 16. A, B, D, †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site; A, close-up of the abdominal region of MGP-PD 26275 showing the embryo; the anterior region of the body lies on the lower portion of the photo; B, reconstruction; D, detail of the vertebral column of the embryo indicated in B with a dotted rectangle. C, dissected specimen of Potamotrygon tigrina (IUWP 7361) showing the position of the left uterus, just next to the stomach. Abbreviations: int, intestine; liv, liver; lu, left uterus; na, neural arches; ru, right uterus; st, stomach; vc, vertebral centra. Scale bars: A, B = 10 mm; D = 1 mm." figureDoi="http://doi.org/10.5281/zenodo.10883311" httpUri="https://zenodo.org/record/10883311/files/figure.png" pageId="17" pageNumber="1205">Fig. 16C</figureCitation>
; but see also Spieler
<emphasis id="B935EA922264B04E273FEA17FCA5FCBE" box="[845,892,859,882]" italics="true" pageId="17" pageNumber="1205">et al</emphasis>
. 2013, fig. 6), and in narcinids in particular (see Nair &amp; Soundararajan 1973, fig. 1;
<bibRefCitation id="EFD04B712264B04E20E0EA37FAEDFC5E" author="Devadoss, P." box="[1170,1332,891,914]" pageId="17" pageNumber="1205" pagination="271 - 283" refId="ref21237" refString="Devadoss, P. 1998. Observations on the breeding and development in some batoid fishes. Indian Journal of Fisheries, 45 (3), 271 - 283." type="journal article" year="1998">Devadoss 1998</bibRefCitation>
, fig. 5). The embryo lies externally to the stomach (whose outline is clearly delimited by its contents of larger foraminifera; see
<figureCitation id="137A2A052264B04E273FEA96FC12FC3D" box="[845,971,986,1009]" captionStart="Figure 16" captionStartId="14.[159,227,1045,1066]" captionTargetBox="[173,1453,183,1013]" captionTargetId="figure-382@14.[173,1453,183,1013]" captionTargetPageId="14" captionText="Figure 16. A, B, D, †Titanonarke molini (Jaekel, 1894) from the Eocene Monte Postale site; A, close-up of the abdominal region of MGP-PD 26275 showing the embryo; the anterior region of the body lies on the lower portion of the photo; B, reconstruction; D, detail of the vertebral column of the embryo indicated in B with a dotted rectangle. C, dissected specimen of Potamotrygon tigrina (IUWP 7361) showing the position of the left uterus, just next to the stomach. Abbreviations: int, intestine; liv, liver; lu, left uterus; na, neural arches; ru, right uterus; st, stomach; vc, vertebral centra. Scale bars: A, B = 10 mm; D = 1 mm." figureDoi="http://doi.org/10.5281/zenodo.10883311" httpUri="https://zenodo.org/record/10883311/files/figure.png" pageId="17" pageNumber="1205">Fig. 16A, B</figureCitation>
) therefore excluding the hypothesis of a possible ingested prey. The absence of traces of egg case surrounding the embryo suggests that the reproductive mode of †
<taxonomicName id="4C414D032264B04E2705ED76FC0EFB9D" baseAuthorityName="Jaekel" baseAuthorityYear="1894" box="[887,983,1082,1105]" class="Elasmobranchii" family="Narcinidae" genus="Titanonarke" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="17" pageNumber="1205" phylum="Chordata" rank="species" species="molini">
<emphasis id="B935EA922264B04E2705ED76FC0EFB9D" box="[887,983,1082,1105]" italics="true" pageId="17" pageNumber="1205">T. molini</emphasis>
</taxonomicName>
was viviparous (probably yolk-sac), a condition that resembles that of most living batoids (
<bibRefCitation id="EFD04B712264B04E213AED16FC1BFB5D" author="Hamlett, W. C. &amp; Koob, T. J." pageId="17" pageNumber="1205" refId="ref22227" refString="Hamlett, W. C. &amp; Koob, T. J. 1999. Chapter 15. Female reproductive system. Pp. 398 - 443 in W. C. Hamlett (ed.) Sharks, skates and rays. The biology of elasmobranch fishes. Johns Hopkins Press, Baltimore." type="book" year="1999">Hamlett &amp; Koob 1999</bibRefCitation>
;
<bibRefCitation id="EFD04B712264B04E27A3ED36FB4BFB5D" author="Kriwet, J. &amp; Kiessling, W. &amp; Klug, S." box="[977,1170,1146,1169]" pageId="17" pageNumber="1205" pagination="945 - 951" refId="ref23744" refString="Kriwet, J., Kiessling, W. &amp; Klug, S. 2009. Diversification trajectories and evolutionary life-history traits in early sharks and batoids. Proceedings of the Royal Society, Series B, 276, 945 - 951." type="journal article" year="2009">
Kriwet
<emphasis id="B935EA922264B04E2053ED36FB96FB5D" box="[1057,1103,1146,1169]" italics="true" pageId="17" pageNumber="1205">et al</emphasis>
. 2009
</bibRefCitation>
), and narcinids in particular (
<bibRefCitation id="EFD04B712264B04E2726EDD6FB84FB7D" author="Hoar, W. S. &amp; Randall, D. J." box="[852,1117,1178,1201]" pageId="17" pageNumber="1205" refId="ref22964" refString="Hoar, W. S. &amp; Randall, D. J. 1988. Fish physiology. Volume XI. The physiology of developing fish - Part B Viviparity and posthatching juveniles. Academic Press, San Diego, 436 pp." type="book" year="1988">Hoar &amp; Randall 1988</bibRefCitation>
;
<bibRefCitation id="EFD04B712264B04E2005EDD6FADEFB7D" author="Bruton, M. N." box="[1143,1287,1178,1201]" pageId="17" pageNumber="1205" refId="ref18921" refString="Bruton, M. N. 1990. Alternative life-history styles of fishes. Developments in environmental biology of fishes 10. Kluwer Academic Publishers, Dordrecht, 327 pp." type="book" year="1990">Bruton 1990</bibRefCitation>
; Rincon 1997; McEachran &amp; Carvalho 2002;
<bibRefCitation id="EFD04B712264B04E20FDEDF6FAEBFB1D" author="Last, P. &amp; White, W. &amp; Carvalho, M. R. de &amp; Stehmann, M. &amp; Naylor, G." box="[1167,1330,1210,1233]" pageId="17" pageNumber="1205" refId="ref23787" refString="Last, P., White, W., Carvalho, M. R. de, S´eret, B., Stehmann, M. &amp; Naylor, G. 2016. Rays of the world. CSIRO Publishing, Clayton North, 800 pp." type="book" year="2016">
Last
<emphasis id="B935EA922264B04E20B1EDF6FB28FB1D" box="[1219,1265,1210,1233]" italics="true" pageId="17" pageNumber="1205">et al</emphasis>
. 2016
</bibRefCitation>
), and is considered plesiomorphic in batoids (
<bibRefCitation id="EFD04B712264B04E20A6ED95FA92FB3C" author="Cole, K. S." box="[1236,1355,1241,1264]" pageId="17" pageNumber="1205" refId="ref20629" refString="Cole, K. S. 2010. Reproduction and sexuality in marine fishes: patterns and processes. University California Press, Berkeley, 432 pp." type="book" year="2010">Cole 2010</bibRefCitation>
). Finally, although the general morphology, size and position of the embryo also resemble those already detected in fossil sharks (e.g. Hovestadt &amp; Hovestadt-Euler 2010; Hovestadt
<emphasis id="B935EA922264B04E273FEC15FC5BFABC" box="[845,898,1369,1392]" italics="true" pageId="17" pageNumber="1205">et al</emphasis>
. 2010) and extinct freshwater stingrays (
<bibRefCitation id="EFD04B712264B04E2128EC15FC66FA5C" author="Carvalho, M. R. de &amp; Maisey, J. C. &amp; Grande, L." pageId="17" pageNumber="1205" pagination="1 - 136" refId="ref20349" refString="Carvalho, M. R. de, Maisey, J. C. &amp; Grande, L. 2004. Freshwater stingrays of the Green River Formation of Wyoming (Early Eocene), with the description of a new genus and species and an analysis of its phylogenetic relationships (Chondrichthyes: Myliobatiformes). Bulletin of the American Museum of Natural History, 284, 1 - 136." type="journal article" year="2004">
Carvalho
<emphasis id="B935EA922264B04E273FEC35FCA5FA5C" box="[845,892,1401,1424]" italics="true" pageId="17" pageNumber="1205">et al</emphasis>
. 2004
</bibRefCitation>
), the specimen described herein unquestionably represents to our knowledge the first occurrence of a fossilized embryo
<emphasis id="B935EA922264B04E27A4ECF5FBCFFA1C" box="[982,1046,1465,1488]" italics="true" pageId="17" pageNumber="1205">in situ</emphasis>
in marine batoid fishes.
</paragraph>
<paragraph id="8BFE36802264B04E273FECB8FB01F907" blockId="17.[845,1467,1524,1739]" pageId="17" pageNumber="1205">
<emphasis id="B935EA922264B04E273FECB8FC65F9C7" bold="true" box="[845,956,1524,1547]" pageId="17" pageNumber="1205">Parasites.</emphasis>
The examination of the historical material also has shown that fossilized crustacean isopods are strictly associated with the body of two individuals of both species of †
<taxonomicName id="4C414D032264B04E27C9EF18FBE4F9A7" box="[955,1085,1620,1643]" class="Elasmobranchii" family="Narcinidae" genus="Titanonarke" higherTaxonomySource="GBIF" kingdom="Animalia" order="Torpediniformes" pageId="17" pageNumber="1205" phylum="Chordata" rank="genus">
<emphasis id="B935EA922264B04E27C9EF18FBE4F9A7" box="[955,1085,1620,1643]" italics="true" pageId="17" pageNumber="1205">Titanonarke</emphasis>
</taxonomicName>
. Individual fishes appear to be infested by two to four isopods each, the analysis of which is beyond the scope of this paper and will be provided in a separate study (Robin
<emphasis id="B935EA922264B04E2049EFF8FBB3F907" box="[1083,1130,1716,1739]" italics="true" pageId="17" pageNumber="1205">et al</emphasis>
. in prep.).
</paragraph>
</subSubSection>
</treatment>
</document>