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<mods:title id="0D86AD02CF414F6ED8008873B129E984">The Clavatulidae (Gastropoda, Conoidea) of the Miocene Paratethys Sea with considerations on fossil and extant Clavatulidae genera</mods:title>
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<heading id="D0CA81ABFFA8FF82FFBAFA296904FA5D" bold="true" box="[151,374,1410,1437]" fontSize="11" level="2" pageId="42" pageNumber="43" reason="4">
<emphasis id="B949EAD5FFA8FF82FFBAFA296904FA5D" bold="true" box="[151,374,1410,1437]" pageId="42" pageNumber="43">
<emphasis id="B949EAD5FFA8FF82FFBAFA296947FA5D" bold="true" box="[151,309,1410,1437]" italics="true" pageId="42" pageNumber="43">bimarginata-</emphasis>
clade
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<paragraph id="8B8236C7FFA8FF82FFBAFA606A1CF88D" blockId="42.[151,1437,1483,1869]" pageId="42" pageNumber="43">
Members of this clade have bipartite early teleoconch sculpture with a granulose subsutural cord and suprasutural spiral cord of equal strength with indistinct, bifid beads (
<figureCitation id="13062A42FFA8FF82FC31FA446B1AF9C9" box="[796,872,1519,1545]" captionStart="FIGURE 3" captionStartId="12.[151,250,1619,1643]" captionTargetBox="[151,1436,181,1595]" captionTargetId="figure-26@12.[151,1436,181,1595]" captionTargetPageId="12" captionText="FIGURE 3. Early teleoconch whorls of extant and fossil Clavatulidae species. A. Clavatula regia (Röding, 1798), NHMW 1683 6/28, West Africa. B. Clavatula rubrifasciata (Reeve, 1845), NHMW 10203, Gorée (Senegal). C. Clavatula diadema (Kiener, 1839), NHMW 1884/0010/0023, Gorée (Senegal). D. Clavatula mystica (Reeve, 1843), MNHN-IM-2013-59897, Est de l'ile de Gorée (Senegal). E. Clavatula bimarginata (Lamarck, 1822), MNHN-IM-2013-43369, Dakar (Senegal). F. Clavatula pyramidata (Kiener, 1839), MNHN-IM-2009-24982, Baie de Pointe-Noire (Republique du Congo). G. Clavatula sorini nov. sp., paratype, NHMW 1874/0025/0046, Lăpugiu de Sus (Romania). H. Clavatula romana (Defrance, 1826), NHMW 1999/0098/0004, Modra-Kráľová (Slovakia). I. Clavatula interrupta (Brocchi, 1814), NHMW 1861/0033/0420, Buituri (Romania). J. Clavatula hirmetzli Kovács &amp; Vicián, 2021, NHMW 1868/0001/0069, Baden-Sooss (Austria). K. Clavatula irisae nov. sp., NHMW 2021/0126/0001, holotype, Gainfarn (Austria). L. Clavatula sophiae (Hoernes &amp; Auinger, 1891), lectotype, NHMW 2021/0127/0001, Enzesfeld (Austria). M. Clavatula veronicae (Hoernes &amp;Auinger, 1891), NHMW 1855/0043/0044b, Lăpugiu de Sus (Romania). N. Clavatula santhai Kovács &amp; Vicián, 2021, holotype, HNHM, PAL 2021.20.1, Letkés (Hungary).O. Clavatula camillae (Hoernes &amp;Auinger, 1891), lectotype, NHMW 1949/0005/0042, Lăpugiu de Sus (Romania). P. Megaclavatula amaliae (Hoernes &amp; Auinger, 1891), lectotype, NHMW 1949/0005/0049, Lăpugiu de Sus (Romania). Q. Megaclavatula grunerti nov. sp., paratype, NHMW 2021/0127/0003, Enzesfeld (Austria). R. Megaclavatula laevigata (Eichwald, 1830), NHMW 1853/0038/0035a, Korytnica (Poland). S. Megaclavatula pilleri nov. sp., holotype, NHMW 1949/0005/0051a, Baden-Sooss (Austria). T. Megaclavatula polonica (Pusch, 1837), NHMW 1862/0001/0324, Bad Vöslau (Austria).U. Granulatocincta angelae (Hoernes &amp;Auinger, 1891), paralectotype, NHMW 1949/0005/0099, Möllersdorf (Austria). V. Granulatocincta granulatocincta (Münster in Goldfuss, 1841), NHMW 1846/0028/0009, Enzesfeld (Austria). W. Granulatocincta nataliae (Hoernes &amp; Auinger, 1891), NHMW 1855/0035/0021, holotype, Belgrade/Rakovica (Serbia). X. Granulatocincta schreibersi (Hörnes, 1854), paralectotype, NHMW1997z0178/1133, Bad Vöslau (Austria).Y. Granulatocincta callim nov. sp., paratype, NHMW 2021/0128/0001, Bad Sauerbrunn (Austria). Z. Granulatocincta contorta (Švagrovský, 1958), holotype, SNMB Z 3076, Kuzmice (Slovakia), Aa. Granulatocincta theoderichi nov. sp., paratype, NHMW 2016/0026/0320b, Ritzing (Austria). Ab. Olegia doderleini (Hörnes, 1854), paralectotype, NHMW 1858/0015/0147, Wiesen (Austria). Ac. Olegia mandici nov. sp., paratype, NHMW 1872/0031/0008a, Bekića vrelo, Čremušnica at Donji Kirin (Croatia). Ae. Olegia dorotheae (Hoernes &amp; Auinger, 1891), paralectotype, NHMW 1949/0005/0040, Kleinebersdorf (Austria). Scale bar is 5 mm for all specimens except for Granulatocincta contorta, Olegia doderleini and O. mandici." figureDoi="http://doi.org/10.5281/zenodo.6399835" httpUri="https://zenodo.org/record/6399835/files/figure.png" pageId="42" pageNumber="43">Fig 3F</figureCitation>
,
<figureCitation id="13062A42FFA8FF82FC58FA446BC5F9C9" box="[885,951,1519,1545]" captionStart="FIGURE 9" captionStartId="22.[151,250,1808,1832]" captionTargetBox="[151,1436,777,1785]" captionTargetId="figure-260@22.[151,1436,777,1785]" captionTargetPageId="22" captionText="FIGURE 9. Extant Clavatulidae species. A. Clavatula rubrifasciata (Reeve, 1845), Gonçalo Rosa, Atoll Specimen Shells (Senegal). B. Clavatula rubrifasciata (Reeve, 1845), NHMW 10203, Gorée (Senegal). C. Clavatula mystica (Reeve, 1843), MNHN-IM-2013-59897, Est de l'ile de Gorée (Senegal). D. Clavatula mystica (Reeve, 1843), Gonçalo Rosa, Atoll Specimen Shells (Senegal). E. Clavatula bimarginata (Lamarck, 1822), MNHN-IM-2013-43369, Dakar (Senegal). F. Clavatula bimarginata (Lamarck, 1822), NHMW 2042, Gorée (Senegal).G. Clavatula diadema (Kiener,1839), NHMW 1884/0010/0023, Gorée (Senegal). Specimens C and E have been used for molecular analysis by Nicolas Puillandre (unpublished data)." figureDoi="http://doi.org/10.5281/zenodo.6399849" httpUri="https://zenodo.org/record/6399849/files/figure.png" pageId="42" pageNumber="43">9EF</figureCitation>
). The moderately concave interspace bears prominent, comma-shaped axial riblets formed by growth lines. Spiral threads form a somewhat cancellate pattern especially in the abapical half of the whorls. Aside from
<emphasis id="B949EAD5FFA8FF82FC0DF99C6BE5F991" box="[800,919,1591,1617]" italics="true" pageId="42" pageNumber="43">
<taxonomicName id="4C3D4D44FFA8FF82FC09F99C6BE5F991" box="[804,919,1591,1617]" class="Gastropoda" family="Clavatulidae" genus="Clavatula" kingdom="Animalia" order="Neogastropoda" pageId="42" pageNumber="43" phylum="Mollusca" rank="genus">Clavatula</taxonomicName>
</emphasis>
<taxonomicName id="4C3D4D44FFA8FF82FC84F99C6D2AF991" authority="(Hoernes &amp; Auinger, 1891)" baseAuthorityName="Hoernes &amp; Auinger" baseAuthorityYear="1891" box="[937,1368,1591,1617]" class="Gastropoda" family="Turridae" genus="Pleurotoma" kingdom="Animalia" order="Neogastropoda" pageId="42" pageNumber="43" phylum="Mollusca" rank="species" species="veronicae">
<emphasis id="B949EAD5FFA8FF82FC84F99C6C64F990" box="[937,1046,1591,1616]" italics="true" pageId="42" pageNumber="43">veronicae</emphasis>
(
<bibRefCitation id="EFAC4B36FFA8FF82FB0AF99C6D22F991" author="Hoernes, R. &amp; Auinger, M." box="[1063,1360,1591,1617]" pageId="42" pageNumber="43" pagination="283 - 382" refId="ref135318" refString="Hoernes, R. &amp; Auinger, M. (1891) Die Gasteropoden der Meeres-Ablagerungen der ersten und zweiten Miocanen Mediterran- Stufe in der Osterreichisch-Ungarischen Monarchie. Abhandlungen der kaiserlich-koniglichen Geologischen Reichsanstalt, 12, 283 - 382, pls 37 - 50. [https: // opac. geologie. ac. at / ais 312 / dokumente / Hoernes % 20 und % 20 Auinger % 20 _ 1879 _ Gasteropoden. pdf]" type="journal article" year="1891">Hoernes &amp; Auinger, 1891</bibRefCitation>
)
</taxonomicName>
, from the middle Miocene of the Central Paratethys Sea, we place here also
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<taxonomicName id="4C3D4D44FFA8FF82FC94F9F06C5EF9B5" box="[953,1068,1627,1653]" class="Gastropoda" family="Clavatulidae" genus="Clavatula" kingdom="Animalia" order="Neogastropoda" pageId="42" pageNumber="43" phylum="Mollusca" rank="genus">Clavatula</taxonomicName>
</emphasis>
<emphasis id="B949EAD5FFA8FF82FB13F9F06CCDF9B5" box="[1086,1215,1627,1653]" italics="true" pageId="42" pageNumber="43">neogradata</emphasis>
<bibRefCitation id="EFAC4B36FFA8FF82FBEAF9F06D2CF9B5" author="Glibert, M." box="[1223,1374,1626,1653]" pageId="42" pageNumber="43" pagination="1 - 75" refId="ref133027" refString="Glibert, M. (1954) Pleurotomes du Miocene de la Belgique et du bassin de la Loire. Memoires de l'Institut Royal des Sciences Naturelles de Belgique, 129, 1 - 75, 7 pls." type="journal article" year="1954">Glibert, 1954</bibRefCitation>
from the Tortonian of
<collectingCountry id="F32A7657FFA8FF82FE79F9D469D2F959" box="[340,416,1663,1689]" name="France" pageId="42" pageNumber="43">France</collectingCountry>
as described by
<bibRefCitation id="EFAC4B36FFA8FF82FD70F9D46B15F959" author="Landau, B. M. &amp; van Dingenen, F. &amp; Ceulemans, L." box="[605,871,1663,1689]" pageId="42" pageNumber="43" pagination="3 - 107" refId="ref137618" refString="Landau, B. M., van Dingenen, F. &amp; Ceulemans, L. (2020) The upper Miocene gastropods of northwestern France, 5. Conoidea. Cainozoic Research, 20, 3 - 107." type="journal article" year="2020">
Landau
<emphasis id="B949EAD5FFA8FF82FD95F9D46A83F959" box="[696,753,1663,1689]" italics="true" pageId="42" pageNumber="43">et al.</emphasis>
(2020: 84
</bibRefCitation>
, pl. 74). The
<emphasis id="B949EAD5FFA8FF82FCD2F9D46CE0F959" box="[1023,1170,1663,1689]" italics="true" pageId="42" pageNumber="43">bimarginata-</emphasis>
clade differs from other bipartite groups:
<taxonomicName id="4C3D4D44FFA8FF82FE78F90869B1F97D" box="[341,451,1699,1725]" class="Gastropoda" family="Clavatulidae" genus="Clavatula" kingdom="Animalia" order="Neogastropoda" pageId="42" pageNumber="43" phylum="Mollusca" rank="genus">
<emphasis id="B949EAD5FFA8FF82FE78F90869B1F97D" box="[341,451,1699,1725]" italics="true" pageId="42" pageNumber="43">Clavatula</emphasis>
</taxonomicName>
(
<emphasis id="B949EAD5FFA8FF82FEFEF908699DF97C" box="[467,495,1699,1724]" italics="true" pageId="42" pageNumber="43">s.s</emphasis>
.) in being more slender fusiform and in having the adsutural cords of equal strength; in
<taxonomicName id="4C3D4D44FFA8FF82FF9BF96C6956F921" box="[182,292,1735,1761]" class="Gastropoda" family="Clavatulidae" genus="Clavatula" kingdom="Animalia" order="Neogastropoda" pageId="42" pageNumber="43" phylum="Mollusca" rank="genus">
<emphasis id="B949EAD5FFA8FF82FF9BF96C6956F921" box="[182,292,1735,1761]" italics="true" pageId="42" pageNumber="43">Clavatula</emphasis>
</taxonomicName>
(
<emphasis id="B949EAD5FFA8FF82FE1AF96C6921F920" box="[311,339,1735,1760]" italics="true" pageId="42" pageNumber="43">s.s</emphasis>
.) the subsutural cord is weak, smooth, and the suprasutural cord is prominent with large beads. The
<taxonomicName id="4C3D4D44FFA8FF82FFFEF9406933F8C5" box="[211,321,1771,1797]" class="Gastropoda" family="Clavatulidae" genus="Clavatula" kingdom="Animalia" order="Neogastropoda" pageId="42" pageNumber="43" phylum="Mollusca" rank="genus">
<emphasis id="B949EAD5FFA8FF82FFFEF9406933F8C5" box="[211,321,1771,1797]" italics="true" pageId="42" pageNumber="43">Clavatula</emphasis>
</taxonomicName>
<taxonomicName id="4C3D4D44FFA8FF82FE7CF94069E3F8C5" baseAuthorityName="Hoernes &amp; Auinger" baseAuthorityYear="1891" box="[337,401,1771,1797]" class="Gastropoda" family="Clavatulidae" genus="Clavatula" kingdom="Animalia" order="Neogastropoda" pageId="42" pageNumber="43" phylum="Mollusca" rank="species" species="juliae">
<emphasis id="B949EAD5FFA8FF82FE7CF94069E3F8C5" box="[337,401,1771,1797]" italics="true" pageId="42" pageNumber="43">juliae</emphasis>
</taxonomicName>
group is immediately separated by its broader profile and strongly constricted base and long siphonal canal. The
<taxonomicName id="4C3D4D44FFA8FF82FEA8F8A46981F8E9" box="[389,499,1807,1833]" class="Gastropoda" family="Clavatulidae" genus="Clavatula" kingdom="Animalia" order="Neogastropoda" pageId="42" pageNumber="43" phylum="Mollusca" rank="genus">
<emphasis id="B949EAD5FFA8FF82FEA8F8A46981F8E9" box="[389,499,1807,1833]" italics="true" pageId="42" pageNumber="43">Clavatula</emphasis>
</taxonomicName>
<taxonomicName id="4C3D4D44FFA8FF82FD2FF8A46A10F8E9" baseAuthorityName="Hoernes R. &amp; Auinger" baseAuthorityYear="1891" box="[514,610,1807,1833]" class="Gastropoda" family="Clavatulidae" genus="Clavatula" kingdom="Animalia" order="Neogastropoda" pageId="42" pageNumber="43" phylum="Mollusca" rank="species" species="camillae">
<emphasis id="B949EAD5FFA8FF82FD2FF8A46A10F8E9" box="[514,610,1807,1833]" italics="true" pageId="42" pageNumber="43">camillae</emphasis>
</taxonomicName>
group also has a slender fusiform profile, but differs in having a smooth concave mid-portion without axial riblets.
</paragraph>
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