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<mods:title id="69AA72047B2417878070DAF64D858749">New Cenozoic Muricidae (Mollusca: Gastropoda) from Europe</mods:title>
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<mods:affiliation id="D7E9C399ED4F532994E9ACDF4659C9C0">Muséum national dhistoire naturelle, Département Origines et Évolution (CR 2 P - MNHN, CNRS, UPMC, Sorbonne Université), 57 rue Cuvier, 75231 Paris cedex 05 (France) didier. merle @ mnhn. fr (corresponding author) jean-michel. pacaud @ mnhn. fr dnledon @ yahoo. fr</mods:affiliation>
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<mods:date id="66DFC1196EDE96D774EC898E1657B92A">2024</mods:date>
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<taxonomicName id="4C274D4FFFFAE41AFC7AFC40FB59FC03" ID-CoL="CN5M7" authorityName="Merle &amp; Pacaud &amp; Ledon &amp; Goret" authorityYear="2024" box="[962,1240,918,944]" class="Gastropoda" family="Muricidae" genus="Timbellus" kingdom="Animalia" order="Neogastropoda" pageId="28" pageNumber="521" phylum="Mollusca" rank="species" species="submicropterus" status="sp. nov.">
<emphasis id="B953EADEFFFAE41AFC7AFC40FB59FC03" bold="true" box="[962,1240,918,944]" italics="true" pageId="28" pageNumber="521">Timbellus submicropterus</emphasis>
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(
<figureCitation id="131C2A49FFFAE41AFC49FC63FBBEFC7C" box="[1009,1087,949,975]" captionStart="FIG" captionStartId="30.[132,143,1872,1889]" captionTargetBox="[147,1440,260,1687]" captionTargetId="figure-206@30.[700,1256,828,1396]" captionTargetPageId="30" captionText="FIG. 16. — Spiral cords and internal denticles of the outer lip in Timbellus submicropterus n. sp. (A-H, K) and T. micropterus (Deshayes, 1835) (I, J, L): A, B, holotype, MNHN.F.A24111 (MNHN coll.), Fresville, Manche, Normandy, France, middle Eocene (Lutetian); C, D, paratype, MNHN.F.A24126 (Munier-Chalmas coll.), Gourbesville, Manche, middle Eocene (Lutetian); E, F, paratype, UCBL-EM 33438 (Bourdot coll.), Fresville; G, H, MNHN.F.A90557 (Brébion coll.), Saint-Aignan- Grand-Lieu (Pierre-Aiguë), Loire-Atlantique, Loire Basin, France, middle Eocene (Bartonian); I, J, MNHN.F.A32522 (Faullummel coll.), Le Limon, Seine-et-Marne, Paris Basin, France,middle Eocene (Bartonian);K, paratype (protoconch), MNHN.F.A90556 (MNHN coll.), Fresville; L, MNHN.F.A90558 (Faullummel coll., protoconch), Chavençon (Carrière Tuyttens), Oise, Paris Basin, France, middle Eocene (Bartonian). Scale bars: A-J, 10 mm; K, L, 0.5 mm. Credits: A-D, I, J, P. Loubry (MNHN/CNRS); E, F, E. Robert (UCBL); G, H, K, L, L. Cazes (MNHN/CNRS)." figureDoi="http://doi.org/10.5281/zenodo.13985800" httpUri="https://zenodo.org/record/13985800/files/figure.png" pageId="28" pageNumber="521">Figs 16</figureCitation>
A-H, K; 27A-D)
</paragraph>
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<taxonomicName id="4C274D4FFFFAE41AFC95FB93FBDBFBEF" ID-CoL="44MK4" authorityName="Lamarck" authorityYear="1803" box="[813,1114,1092,1116]" class="Gastropoda" family="Muricidae" genus="Murex" kingdom="Animalia" order="Neogastropoda" pageId="28" pageNumber="521" phylum="Mollusca" rank="species" species="tricarinatus" subGenus="Pterymurex">
<emphasis id="B953EADEFFFAE41AFC95FB93FCEBFBEF" box="[813,874,1093,1116]" italics="true" pageId="28" pageNumber="521">Murex</emphasis>
(
<emphasis id="B953EADEFFFAE41AFCCFFB93FC60FBEF" box="[887,993,1093,1116]" italics="true" pageId="28" pageNumber="521">Pterymurex</emphasis>
)
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<bibRefCitation id="EFB64B3DFFFAE41AFBD7FB92FA2AFBEF" author="COSSMANN M. &amp; PISSARRO G." box="[1135,1451,1092,1116]" pageId="28" pageNumber="521" pagination="61 - 140" refId="ref43809" refString="COSSMANN M. &amp; PISSARRO G. 1901. - Faune eocenique du Cotentin. 2 eme article. Bulletin de la Societe Geologique de Normandie 20: 11 - 90. [Also published as Cossmann M. &amp; Pissarro G. 1901. - Faune eocenique du Cotentin (Mollusques), Tome 1, Fascicule 2, Imprimerie du Journal Le Havre, Le Havre, 61 - 140]." type="book chapter" year="1901">Cossmann &amp; Pissarro 1901: 128</bibRefCitation>
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, pl. 14, fig. 1 [
<emphasis id="B953EADEFFFAE41AFC0BFB89FC56FBC5" box="[947,983,1119,1142]" italics="true" pageId="28" pageNumber="521">non</emphasis>
<bibRefCitation id="EFB64B3DFFFAE41AFC65FB89FBF0FBC4" author="LAMARCK J. - B." box="[989,1137,1119,1143]" pageId="28" pageNumber="521" pagination="217 - 227" refId="ref45566" refString="LAMARCK J. - B. 1803. - Memoires sur les fossiles des environs de Paris (suite 1). Annales du Museum d'Histoire naturelle 2 (9): 217 - 227." type="journal article" year="1803">Lamarck, 1803</bibRefCitation>
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<taxonomicName id="4C274D4FFFFAE41AFC95FB4EFC7CFB1C" ID-CoL="86YQB" baseAuthorityName="Deshayes" baseAuthorityYear="1835" box="[813,1021,1176,1200]" class="Gastropoda" family="Muricidae" genus="Timbellus" kingdom="Animalia" order="Neogastropoda" pageId="28" pageNumber="521" phylum="Mollusca" rank="species" species="micropterus">
<emphasis id="B953EADEFFFAE41AFC95FB4EFC7CFB1C" box="[813,1021,1176,1200]" italics="true" pageId="28" pageNumber="521">Timbellus micropterus</emphasis>
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<bibRefCitation id="EFB64B3DFFFAE41AFBA1FB4EFB79FB03" author="MERLE D. &amp; GARRIGUES B. &amp; POINTIER J. - P." box="[1049,1272,1176,1200]" pageId="28" pageNumber="521" refId="ref47234" refString="MERLE D., GARRIGUES B. &amp; POINTIER J. - P. 2011. - Fossil &amp; Recent Muricidae of the World. Part Muricinae. ConchBooks, Hackenheim, 652 p." type="book" year="2011">
Merle
<emphasis id="B953EADEFFFAE41AFBE1FB4EFB08FB03" box="[1113,1161,1176,1200]" italics="true" pageId="28" pageNumber="521">et al.</emphasis>
2011: 448
</bibRefCitation>
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, pl. 102, fig. 4a, b only [
<emphasis id="B953EADEFFFAE41AFCDEFB65FC0BFB79" box="[870,906,1203,1226]" italics="true" pageId="28" pageNumber="521">non</emphasis>
<bibRefCitation id="EFB64B3DFFFAE41AFC28FB64FBABFB79" author="DESHAYES G. - P." box="[912,1066,1202,1226]" pageId="28" pageNumber="521" pagination="101 - 107" refId="ref44001" refString="DESHAYES G. - P. 1865. - Description des Animaux sans vertebres decouverts dans le bassin de Paris pour servir de supplement a la description des coquilles fossiles des environs de Paris, comprenant une revue generale de toutes les especes actuellement connues. Tome 3. Bailliere, Paris, Livraisons 45 - 46, p. 201 - 288; Atlas 2, pl. 86 - 93 [2 January]; Livraisons 47 - 48, p. 289 - 424, Atlas 2, pl. 94 - 100 [10 June]; Livraisons 49 - 50, p. 425 - 668; Atlas 2, pl. 101 - 107 [15 December]." type="journal article" year="1865">Deshayes, 1865</bibRefCitation>
].
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. —
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.
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, Manche, Fresville;
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Calcaire
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; middle Eocene (uppermost Lutetian);
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.F.A24111 (
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coll.), H:
<quantity id="4CDF9B29FFFAE41AFB52FACAFAC5FA87" box="[1258,1348,1308,1332]" metricMagnitude="-2" metricUnit="m" metricValue="2.36" pageId="28" pageNumber="521" unit="mm" value="23.6">23.6 mm</quantity>
(
<figureCitation id="131C2A49FFFAE41AFAEAFACAFCBEFAFD" captionStart="FIG" captionStartId="30.[132,143,1872,1889]" captionTargetBox="[147,1440,260,1687]" captionTargetId="figure-206@30.[700,1256,828,1396]" captionTargetPageId="30" captionText="FIG. 16. — Spiral cords and internal denticles of the outer lip in Timbellus submicropterus n. sp. (A-H, K) and T. micropterus (Deshayes, 1835) (I, J, L): A, B, holotype, MNHN.F.A24111 (MNHN coll.), Fresville, Manche, Normandy, France, middle Eocene (Lutetian); C, D, paratype, MNHN.F.A24126 (Munier-Chalmas coll.), Gourbesville, Manche, middle Eocene (Lutetian); E, F, paratype, UCBL-EM 33438 (Bourdot coll.), Fresville; G, H, MNHN.F.A90557 (Brébion coll.), Saint-Aignan- Grand-Lieu (Pierre-Aiguë), Loire-Atlantique, Loire Basin, France, middle Eocene (Bartonian); I, J, MNHN.F.A32522 (Faullummel coll.), Le Limon, Seine-et-Marne, Paris Basin, France,middle Eocene (Bartonian);K, paratype (protoconch), MNHN.F.A90556 (MNHN coll.), Fresville; L, MNHN.F.A90558 (Faullummel coll., protoconch), Chavençon (Carrière Tuyttens), Oise, Paris Basin, France, middle Eocene (Bartonian). Scale bars: A-J, 10 mm; K, L, 0.5 mm. Credits: A-D, I, J, P. Loubry (MNHN/CNRS); E, F, E. Robert (UCBL); G, H, K, L, L. Cazes (MNHN/CNRS)." figureDoi="http://doi.org/10.5281/zenodo.13985800" httpUri="https://zenodo.org/record/13985800/files/figure.png" pageId="28" pageNumber="521">Figs 16A, B</figureCitation>
;
<figureCitation id="131C2A49FFFAE41AFCF2FAE1FCF9FAFC" box="[842,888,1335,1359]" captionStart="FIG" captionStartId="46.[132,143,1606,1623]" captionTargetBox="[132,1455,215,1546]" captionTargetId="figure-366@46.[699,1260,794,1359]" captionTargetPageId="46" captionText="FIG. 27. — New species of Timbellus De Gregorio, 1885, Pterynotus Swainson, 1833 (Pterymarchia Houart, 1995), Textiliomurex Merle, 2011, Ponderia Houart, 1986 and Flexopteron Shuto, 1969: A-C, Timbellus submicropterus n. sp.: A, holotype, MNHN.F.A24111 (MNHN coll.), Fresville, France, Lutetian; B, paratype, MNHN.F.A24126 (Munier-Chalmas coll.), Gourbesville, France, Lutetian; C, paratype, UCBL-EM 33438 (Bourdot coll.), Fresville; D, T. cf. submicropterus n. sp., MNHN.F.A90557 (Brébion coll.), Saint-Aignan-Grand-Lieu (Pierre-Aiguë), France, Bartonian; E, F, Pterynotus (Pterymarchia) gaasensis n. sp.: E, holotype, MNHN.F.A32492 (Merle coll.), Gaas (Espibos), France, Rupelian; F, paratype, MNHN.F.A90554 (Merle coll.), Gaas (Espibos); G, H, P. (P.) pelouatensis n. sp.: G, holotype, MNHN.F.J09945, Saucats (Peloua), France, Burdigalian; H, paratype, MNHN.F.J17632, Saucats (Peloua); I, J, Textiliomurex chodmonensis n. sp.: I, holotype, MNHN.F.A81144 (Pacaud coll.), Chaumont-en-Vexin (Carrière Darcy), France, Lutetian; J, paratype, MNHN.F.A81145 (Pons coll.), Chaumont-en- Vexin (Carrière Darcy); K, L, Ponderia remyi n. sp.: K, holotype, MNHN.F.A90548 (Remy coll.), Fresville, Lutetian; L, paratype, MNHN.F.A90555, Fresville (Remy coll.); M, N, Flexopteron constantinense n. sp.: M, paratype, MNHN.F.A77761 (MNHN coll.), Carentan-les-Marais (Cauvin), France, Priabonian; N, holotype, MNHN.F.A77760 (MNHN coll.), Carentan-les-Marais (Cauvin). Scale bars: A-D, I-L, 10 mm; E-H, N, 5 mm; M, 3 mm. Credits: A, B, D-F, N, P. Loubry (MNHN/ CNRS); C, E. Robert (UCBL)." figureDoi="http://doi.org/10.5281/zenodo.13985831" httpUri="https://zenodo.org/record/13985831/files/figure.png" pageId="28" pageNumber="521">27A</figureCitation>
).
</materialsCitation>
</paragraph>
<paragraph id="8B9836CCFFFAE41AFC95FA87FB8CF990" blockId="28.[811,1457,1255,1571]" pageId="28" pageNumber="521">
<materialsCitation id="3B4F3C91FFFAE41AFC95FA87FB92FA2D" ID-GBIF-Occurrence="4958671426" collectorName="Bourdot" country="France" location="France" pageId="28" pageNumber="521" specimenCode="UCBL-EM 33438" specimenCount="1" typeStatus="paratype">
<emphasis id="B953EADEFFFAE41AFC95FA87FC5DFADA" bold="true" box="[813,988,1361,1385]" pageId="28" pageNumber="521">
<typeStatus id="549C886EFFFAE41AFC95FA87FC0EFADA" box="[813,911,1361,1385]" pageId="28" pageNumber="521" type="paratype">Paratypes</typeStatus>
.
<collectingCountry id="F330765CFFFAE41AFC22FA87FC5DFADA" box="[922,988,1361,1385]" name="France" pageId="28" pageNumber="521">France</collectingCountry>
</emphasis>
• 1 spm; same as for the holotype;
<specimenCode id="DB819EB7FFFAE41AFAF9FA87FCECFA37" collectionCode="UCBL-EM" pageId="28" pageNumber="521">UCBL-EM 33438</specimenCode>
(
<collectorName id="26D2531AFFFAE41AFCC4FABAFC4EFA37" box="[892,975,1388,1412]" pageId="28" pageNumber="521">Bourdot</collectorName>
coll., figured specimen of
<bibRefCitation id="EFB64B3DFFFAE41AFB61FABAFCDEFA2D" author="COSSMANN M. &amp; PISSARRO G." pageId="28" pageNumber="521" pagination="61 - 140" refId="ref43809" refString="COSSMANN M. &amp; PISSARRO G. 1901. - Faune eocenique du Cotentin. 2 eme article. Bulletin de la Societe Geologique de Normandie 20: 11 - 90. [Also published as Cossmann M. &amp; Pissarro G. 1901. - Faune eocenique du Cotentin (Mollusques), Tome 1, Fascicule 2, Imprimerie du Journal Le Havre, Le Havre, 61 - 140]." type="book chapter" year="1901">Cossmann &amp; Pissarro 1901</bibRefCitation>
)(
<figureCitation id="131C2A49FFFAE41AFCC9FA50FC5BFA2D" box="[881,986,1414,1438]" captionStart="FIG" captionStartId="30.[132,143,1872,1889]" captionTargetBox="[147,1440,260,1687]" captionTargetId="figure-206@30.[700,1256,828,1396]" captionTargetPageId="30" captionText="FIG. 16. — Spiral cords and internal denticles of the outer lip in Timbellus submicropterus n. sp. (A-H, K) and T. micropterus (Deshayes, 1835) (I, J, L): A, B, holotype, MNHN.F.A24111 (MNHN coll.), Fresville, Manche, Normandy, France, middle Eocene (Lutetian); C, D, paratype, MNHN.F.A24126 (Munier-Chalmas coll.), Gourbesville, Manche, middle Eocene (Lutetian); E, F, paratype, UCBL-EM 33438 (Bourdot coll.), Fresville; G, H, MNHN.F.A90557 (Brébion coll.), Saint-Aignan- Grand-Lieu (Pierre-Aiguë), Loire-Atlantique, Loire Basin, France, middle Eocene (Bartonian); I, J, MNHN.F.A32522 (Faullummel coll.), Le Limon, Seine-et-Marne, Paris Basin, France,middle Eocene (Bartonian);K, paratype (protoconch), MNHN.F.A90556 (MNHN coll.), Fresville; L, MNHN.F.A90558 (Faullummel coll., protoconch), Chavençon (Carrière Tuyttens), Oise, Paris Basin, France, middle Eocene (Bartonian). Scale bars: A-J, 10 mm; K, L, 0.5 mm. Credits: A-D, I, J, P. Loubry (MNHN/CNRS); E, F, E. Robert (UCBL); G, H, K, L, L. Cazes (MNHN/CNRS)." figureDoi="http://doi.org/10.5281/zenodo.13985800" httpUri="https://zenodo.org/record/13985800/files/figure.png" pageId="28" pageNumber="521">Figs 16E, F</figureCitation>
;
<figureCitation id="131C2A49FFFAE41AFC58FA50FB8DFA2D" box="[992,1036,1414,1438]" captionStart="FIG" captionStartId="46.[132,143,1606,1623]" captionTargetBox="[132,1455,215,1546]" captionTargetId="figure-366@46.[699,1260,794,1359]" captionTargetPageId="46" captionText="FIG. 27. — New species of Timbellus De Gregorio, 1885, Pterynotus Swainson, 1833 (Pterymarchia Houart, 1995), Textiliomurex Merle, 2011, Ponderia Houart, 1986 and Flexopteron Shuto, 1969: A-C, Timbellus submicropterus n. sp.: A, holotype, MNHN.F.A24111 (MNHN coll.), Fresville, France, Lutetian; B, paratype, MNHN.F.A24126 (Munier-Chalmas coll.), Gourbesville, France, Lutetian; C, paratype, UCBL-EM 33438 (Bourdot coll.), Fresville; D, T. cf. submicropterus n. sp., MNHN.F.A90557 (Brébion coll.), Saint-Aignan-Grand-Lieu (Pierre-Aiguë), France, Bartonian; E, F, Pterynotus (Pterymarchia) gaasensis n. sp.: E, holotype, MNHN.F.A32492 (Merle coll.), Gaas (Espibos), France, Rupelian; F, paratype, MNHN.F.A90554 (Merle coll.), Gaas (Espibos); G, H, P. (P.) pelouatensis n. sp.: G, holotype, MNHN.F.J09945, Saucats (Peloua), France, Burdigalian; H, paratype, MNHN.F.J17632, Saucats (Peloua); I, J, Textiliomurex chodmonensis n. sp.: I, holotype, MNHN.F.A81144 (Pacaud coll.), Chaumont-en-Vexin (Carrière Darcy), France, Lutetian; J, paratype, MNHN.F.A81145 (Pons coll.), Chaumont-en- Vexin (Carrière Darcy); K, L, Ponderia remyi n. sp.: K, holotype, MNHN.F.A90548 (Remy coll.), Fresville, Lutetian; L, paratype, MNHN.F.A90555, Fresville (Remy coll.); M, N, Flexopteron constantinense n. sp.: M, paratype, MNHN.F.A77761 (MNHN coll.), Carentan-les-Marais (Cauvin), France, Priabonian; N, holotype, MNHN.F.A77760 (MNHN coll.), Carentan-les-Marais (Cauvin). Scale bars: A-D, I-L, 10 mm; E-H, N, 5 mm; M, 3 mm. Credits: A, B, D-F, N, P. Loubry (MNHN/ CNRS); C, E. Robert (UCBL)." figureDoi="http://doi.org/10.5281/zenodo.13985831" httpUri="https://zenodo.org/record/13985831/files/figure.png" pageId="28" pageNumber="521">27C</figureCitation>
)
</materialsCitation>
<materialsCitation id="3B4F3C91FFFAE41AFBAFFA51FC4EFA0A" ID-GBIF-Occurrence="4958671418" collectorName="Bourdot" country="France" location="France" pageId="28" pageNumber="521" specimenCode="UCBL-EM 33438" specimenCount="1" typeStatus="paratype">
• 1 spm; idem;
<collectionCode id="ED36AE09FFFAE41AFB26FA51FB71FA2D" box="[1182,1264,1415,1438]" country="France" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34988" lsid="urn:lsid:biocol.org:col:34988" name="Museum National d'Histoire Naturelle" pageId="28" pageNumber="521">MNHN</collectionCode>
.F.A90556 (
<collectionCode id="ED36AE09FFFAE41AFAE5FA50FA2EFA2D" box="[1373,1455,1414,1438]" country="France" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34988" lsid="urn:lsid:biocol.org:col:34988" name="Museum National d'Histoire Naturelle" pageId="28" pageNumber="521">MNHN</collectionCode>
coll.) (
<figureCitation id="131C2A49FFFAE41AFCD7FA77FC49FA0A" box="[879,968,1441,1465]" captionStart="FIG" captionStartId="30.[132,143,1872,1889]" captionTargetBox="[147,1440,260,1687]" captionTargetId="figure-206@30.[700,1256,828,1396]" captionTargetPageId="30" captionText="FIG. 16. — Spiral cords and internal denticles of the outer lip in Timbellus submicropterus n. sp. (A-H, K) and T. micropterus (Deshayes, 1835) (I, J, L): A, B, holotype, MNHN.F.A24111 (MNHN coll.), Fresville, Manche, Normandy, France, middle Eocene (Lutetian); C, D, paratype, MNHN.F.A24126 (Munier-Chalmas coll.), Gourbesville, Manche, middle Eocene (Lutetian); E, F, paratype, UCBL-EM 33438 (Bourdot coll.), Fresville; G, H, MNHN.F.A90557 (Brébion coll.), Saint-Aignan- Grand-Lieu (Pierre-Aiguë), Loire-Atlantique, Loire Basin, France, middle Eocene (Bartonian); I, J, MNHN.F.A32522 (Faullummel coll.), Le Limon, Seine-et-Marne, Paris Basin, France,middle Eocene (Bartonian);K, paratype (protoconch), MNHN.F.A90556 (MNHN coll.), Fresville; L, MNHN.F.A90558 (Faullummel coll., protoconch), Chavençon (Carrière Tuyttens), Oise, Paris Basin, France, middle Eocene (Bartonian). Scale bars: A-J, 10 mm; K, L, 0.5 mm. Credits: A-D, I, J, P. Loubry (MNHN/CNRS); E, F, E. Robert (UCBL); G, H, K, L, L. Cazes (MNHN/CNRS)." figureDoi="http://doi.org/10.5281/zenodo.13985800" httpUri="https://zenodo.org/record/13985800/files/figure.png" pageId="28" pageNumber="521">Fig. 16K</figureCitation>
)
</materialsCitation>
<materialsCitation id="3B4F3C91FFFAE41AFC51FA77FCE3FA60" ID-GBIF-Occurrence="4958671379" collectorName="Bourdot" country="France" pageId="28" pageNumber="521" specimenCount="1" typeStatus="paratype">
1 spm; idem;
<collectionCode id="ED36AE09FFFAE41AFBC0FA77FB4DFA0B" box="[1144,1228,1441,1464]" country="France" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34988" lsid="urn:lsid:biocol.org:col:34988" name="Museum National d'Histoire Naturelle" pageId="28" pageNumber="521">MNHN</collectionCode>
.F.B63700 (
<collectorName id="26D2531AFFFAE41AFAF8FA77FA2EFA0A" box="[1344,1455,1441,1465]" pageId="28" pageNumber="521">de Morgan</collectorName>
coll.)
</materialsCitation>
<materialsCitation id="3B4F3C91FFFAE41AFCC6FA6DFB81F990" ID-GBIF-Occurrence="4958671370" collectorName="Manche &amp; Gourbesville" country="France" location="Normandy" pageId="28" pageNumber="521" specimenCount="1" stateProvince="Haute-Normandie" typeStatus="paratype">
1 spm;
<collectingRegion id="49E3F82EFFFAE41AFC76FA6DFBBFFA60" box="[974,1086,1467,1491]" country="France" name="Haute-Normandie" pageId="28" pageNumber="521">Normandy</collectingRegion>
,
<collectorName id="26D2531AFFFAE41AFBF1FA6DFB1EFA60" box="[1097,1183,1467,1491]" pageId="28" pageNumber="521">Manche</collectorName>
,
<collectorName id="26D2531AFFFAE41AFB13FA6DFAB1FA60" box="[1195,1328,1467,1491]" pageId="28" pageNumber="521">Gourbesville</collectorName>
;
<location id="8EF86017FFFAE41AFA84FA6DFC05FA5D" LSID="urn:lsid:plazi:treatment:038E87DAFFFAE41BFC7AFC40FC86F858:8EF86017FFFAE41AFA84FA6DFC05FA5D" country="France" name="Calcaire de Fresville" pageId="28" pageNumber="521">Calcaire de Fresville</location>
;
<location id="8EF86017FFFAE41AFC37FA00FBAAFA5D" LSID="urn:lsid:plazi:treatment:038E87DAFFFAE41BFC7AFC40FC86F858:8EF86017FFFAE41AFC37FA00FBAAFA5D" box="[911,1067,1494,1518]" country="France" name="upper Lutetian" pageId="28" pageNumber="521">upper Lutetian</location>
;
<collectionCode id="ED36AE09FFFAE41AFB8EFA00FB0AFA5E" box="[1078,1163,1494,1517]" country="France" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34988" lsid="urn:lsid:biocol.org:col:34988" name="Museum National d'Histoire Naturelle" pageId="28" pageNumber="521">MNHN</collectionCode>
.F.A24126 (
<collectorName id="26D2531AFFFAE41AFABAFA00FA2EFA5D" box="[1282,1455,1494,1518]" pageId="28" pageNumber="521">Munier-Chalmas</collectorName>
coll.), figured specimen of
<bibRefCitation id="EFB64B3DFFFAE41AFB97FA26FA90F9BB" author="MERLE D. &amp; GARRIGUES B. &amp; POINTIER J. - P." box="[1071,1297,1520,1544]" pageId="28" pageNumber="521" refId="ref47234" refString="MERLE D., GARRIGUES B. &amp; POINTIER J. - P. 2011. - Fossil &amp; Recent Muricidae of the World. Part Muricinae. ConchBooks, Hackenheim, 652 p." type="book" year="2011">
<taxonomicName id="4C274D4FFFFAE41AFB97FA26FBE9F9BB" authorityName="Merle &amp; Pacaud &amp; Ledon &amp; Goret" authorityYear="2024" box="[1071,1128,1520,1544]" class="Gastropoda" family="Muricidae" kingdom="Animalia" order="Neogastropoda" pageId="28" pageNumber="543" phylum="Mollusca" rank="subFamily" status="subfam. nov." subFamily="Merle">Merle</taxonomicName>
<emphasis id="B953EADEFFFAE41AFBD6FA27FB1CF9BB" box="[1134,1181,1520,1544]" italics="true" pageId="28" pageNumber="521">et al.</emphasis>
(2011: 135
</bibRefCitation>
, pl. 102, fig. 4a, b) (
<figureCitation id="131C2A49FFFAE41AFCE8F9DCFC4AF991" box="[848,971,1546,1571]" captionStart="FIG" captionStartId="30.[132,143,1872,1889]" captionTargetBox="[147,1440,260,1687]" captionTargetId="figure-206@30.[700,1256,828,1396]" captionTargetPageId="30" captionText="FIG. 16. — Spiral cords and internal denticles of the outer lip in Timbellus submicropterus n. sp. (A-H, K) and T. micropterus (Deshayes, 1835) (I, J, L): A, B, holotype, MNHN.F.A24111 (MNHN coll.), Fresville, Manche, Normandy, France, middle Eocene (Lutetian); C, D, paratype, MNHN.F.A24126 (Munier-Chalmas coll.), Gourbesville, Manche, middle Eocene (Lutetian); E, F, paratype, UCBL-EM 33438 (Bourdot coll.), Fresville; G, H, MNHN.F.A90557 (Brébion coll.), Saint-Aignan- Grand-Lieu (Pierre-Aiguë), Loire-Atlantique, Loire Basin, France, middle Eocene (Bartonian); I, J, MNHN.F.A32522 (Faullummel coll.), Le Limon, Seine-et-Marne, Paris Basin, France,middle Eocene (Bartonian);K, paratype (protoconch), MNHN.F.A90556 (MNHN coll.), Fresville; L, MNHN.F.A90558 (Faullummel coll., protoconch), Chavençon (Carrière Tuyttens), Oise, Paris Basin, France, middle Eocene (Bartonian). Scale bars: A-J, 10 mm; K, L, 0.5 mm. Credits: A-D, I, J, P. Loubry (MNHN/CNRS); E, F, E. Robert (UCBL); G, H, K, L, L. Cazes (MNHN/CNRS)." figureDoi="http://doi.org/10.5281/zenodo.13985800" httpUri="https://zenodo.org/record/13985800/files/figure.png" pageId="28" pageNumber="521">Figs 16C, D</figureCitation>
;
<figureCitation id="131C2A49FFFAE41AFC6DF9DDFB81F990" box="[981,1024,1547,1571]" captionStart="FIG" captionStartId="46.[132,143,1606,1623]" captionTargetBox="[132,1455,215,1546]" captionTargetId="figure-366@46.[699,1260,794,1359]" captionTargetPageId="46" captionText="FIG. 27. — New species of Timbellus De Gregorio, 1885, Pterynotus Swainson, 1833 (Pterymarchia Houart, 1995), Textiliomurex Merle, 2011, Ponderia Houart, 1986 and Flexopteron Shuto, 1969: A-C, Timbellus submicropterus n. sp.: A, holotype, MNHN.F.A24111 (MNHN coll.), Fresville, France, Lutetian; B, paratype, MNHN.F.A24126 (Munier-Chalmas coll.), Gourbesville, France, Lutetian; C, paratype, UCBL-EM 33438 (Bourdot coll.), Fresville; D, T. cf. submicropterus n. sp., MNHN.F.A90557 (Brébion coll.), Saint-Aignan-Grand-Lieu (Pierre-Aiguë), France, Bartonian; E, F, Pterynotus (Pterymarchia) gaasensis n. sp.: E, holotype, MNHN.F.A32492 (Merle coll.), Gaas (Espibos), France, Rupelian; F, paratype, MNHN.F.A90554 (Merle coll.), Gaas (Espibos); G, H, P. (P.) pelouatensis n. sp.: G, holotype, MNHN.F.J09945, Saucats (Peloua), France, Burdigalian; H, paratype, MNHN.F.J17632, Saucats (Peloua); I, J, Textiliomurex chodmonensis n. sp.: I, holotype, MNHN.F.A81144 (Pacaud coll.), Chaumont-en-Vexin (Carrière Darcy), France, Lutetian; J, paratype, MNHN.F.A81145 (Pons coll.), Chaumont-en- Vexin (Carrière Darcy); K, L, Ponderia remyi n. sp.: K, holotype, MNHN.F.A90548 (Remy coll.), Fresville, Lutetian; L, paratype, MNHN.F.A90555, Fresville (Remy coll.); M, N, Flexopteron constantinense n. sp.: M, paratype, MNHN.F.A77761 (MNHN coll.), Carentan-les-Marais (Cauvin), France, Priabonian; N, holotype, MNHN.F.A77760 (MNHN coll.), Carentan-les-Marais (Cauvin). Scale bars: A-D, I-L, 10 mm; E-H, N, 5 mm; M, 3 mm. Credits: A, B, D-F, N, P. Loubry (MNHN/ CNRS); C, E. Robert (UCBL)." figureDoi="http://doi.org/10.5281/zenodo.13985831" httpUri="https://zenodo.org/record/13985831/files/figure.png" pageId="28" pageNumber="521">27B</figureCitation>
</materialsCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="C33D6547FFFAE41AFC95F997FA31F96D" pageId="28" pageNumber="521" type="description">
<paragraph id="8B9836CCFFFAE41AFC95F997FA31F96D" blockId="28.[813,1457,1601,1758]" pageId="28" pageNumber="521">
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(material referring to
<taxonomicName id="4C274D4FFFFAE41AFB46F997FC1CF9C0" authorityName="Merle &amp; Pacaud &amp; Ledon &amp; Goret" authorityYear="2024" class="Gastropoda" family="Muricidae" genus="Timbellus" isUncertain="true" kingdom="Animalia" order="Neogastropoda" pageId="28" pageNumber="521" phylum="Mollusca" rank="species" species="submicropterus" status="sp. nov.">
<emphasis id="B953EADEFFFAE41AFB46F997FADEF9EA" box="[1278,1375,1601,1625]" italics="true" pageId="28" pageNumber="521">Timbellus</emphasis>
cf.
<emphasis id="B953EADEFFFAE41AFA3FF997FC1CF9C0" italics="true" pageId="28" pageNumber="521">submicropterus</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A26057A5FFFAE41AFC1DF98AFC61F9C7" box="[933,992,1628,1652]" pageId="28" pageNumber="521" rank="species">n. sp.</taxonomicNameLabel>
). —
<materialsCitation id="3B4F3C91FFFAE41AFBA1F98AFBF6F91A" ID-GBIF-Occurrence="4958671347" pageId="28" pageNumber="521">
<emphasis id="B953EADEFFFAE41AFBA1F98AFBDCF9C7" bold="true" box="[1049,1117,1628,1652]" pageId="28" pageNumber="521">
<collectingCountry id="F330765CFFFAE41AFBA1F98AFBDCF9C7" box="[1049,1117,1628,1652]" name="France" pageId="28" pageNumber="521">France</collectingCountry>
</emphasis>
• 3 spm; Loire-Atlantique,
<collectingRegion id="49E3F82EFFFAE41AFAC3F98AFA2EF9C7" box="[1403,1455,1628,1652]" country="France" name="Pays de la Loire" pageId="28" pageNumber="521">Loire</collectingRegion>
Basin, Saffré (Bois Gouët); middle Eocene (lower Bartonian); MNHN.F.A29660 (Pacaud coll.)
</materialsCitation>
<materialsCitation id="3B4F3C91FFFAE41AFB29F947FA31F96D" ID-GBIF-Occurrence="4958671364" pageId="28" pageNumber="521">
2 spm; Saint-Aignan-GrandLieu (Pierre-Aiguë), Loire-Atlantique; idem; MNHN.F.A29662, (Brébion coll.),MNHN.F.A90557 (Brébion coll. (
<figureCitation id="131C2A49FFFAE41AFB42F913FAEFF96E" box="[1274,1390,1733,1758]" captionStart="FIG" captionStartId="30.[132,143,1872,1889]" captionTargetBox="[147,1440,260,1687]" captionTargetId="figure-206@30.[700,1256,828,1396]" captionTargetPageId="30" captionText="FIG. 16. — Spiral cords and internal denticles of the outer lip in Timbellus submicropterus n. sp. (A-H, K) and T. micropterus (Deshayes, 1835) (I, J, L): A, B, holotype, MNHN.F.A24111 (MNHN coll.), Fresville, Manche, Normandy, France, middle Eocene (Lutetian); C, D, paratype, MNHN.F.A24126 (Munier-Chalmas coll.), Gourbesville, Manche, middle Eocene (Lutetian); E, F, paratype, UCBL-EM 33438 (Bourdot coll.), Fresville; G, H, MNHN.F.A90557 (Brébion coll.), Saint-Aignan- Grand-Lieu (Pierre-Aiguë), Loire-Atlantique, Loire Basin, France, middle Eocene (Bartonian); I, J, MNHN.F.A32522 (Faullummel coll.), Le Limon, Seine-et-Marne, Paris Basin, France,middle Eocene (Bartonian);K, paratype (protoconch), MNHN.F.A90556 (MNHN coll.), Fresville; L, MNHN.F.A90558 (Faullummel coll., protoconch), Chavençon (Carrière Tuyttens), Oise, Paris Basin, France, middle Eocene (Bartonian). Scale bars: A-J, 10 mm; K, L, 0.5 mm. Credits: A-D, I, J, P. Loubry (MNHN/CNRS); E, F, E. Robert (UCBL); G, H, K, L, L. Cazes (MNHN/CNRS)." figureDoi="http://doi.org/10.5281/zenodo.13985800" httpUri="https://zenodo.org/record/13985800/files/figure.png" pageId="28" pageNumber="521">Figs 16G, H</figureCitation>
;
<figureCitation id="131C2A49FFFAE41AFACCF910FA22F96D" box="[1396,1443,1734,1758]" captionStart="FIG" captionStartId="46.[132,143,1606,1623]" captionTargetBox="[132,1455,215,1546]" captionTargetId="figure-366@46.[699,1260,794,1359]" captionTargetPageId="46" captionText="FIG. 27. — New species of Timbellus De Gregorio, 1885, Pterynotus Swainson, 1833 (Pterymarchia Houart, 1995), Textiliomurex Merle, 2011, Ponderia Houart, 1986 and Flexopteron Shuto, 1969: A-C, Timbellus submicropterus n. sp.: A, holotype, MNHN.F.A24111 (MNHN coll.), Fresville, France, Lutetian; B, paratype, MNHN.F.A24126 (Munier-Chalmas coll.), Gourbesville, France, Lutetian; C, paratype, UCBL-EM 33438 (Bourdot coll.), Fresville; D, T. cf. submicropterus n. sp., MNHN.F.A90557 (Brébion coll.), Saint-Aignan-Grand-Lieu (Pierre-Aiguë), France, Bartonian; E, F, Pterynotus (Pterymarchia) gaasensis n. sp.: E, holotype, MNHN.F.A32492 (Merle coll.), Gaas (Espibos), France, Rupelian; F, paratype, MNHN.F.A90554 (Merle coll.), Gaas (Espibos); G, H, P. (P.) pelouatensis n. sp.: G, holotype, MNHN.F.J09945, Saucats (Peloua), France, Burdigalian; H, paratype, MNHN.F.J17632, Saucats (Peloua); I, J, Textiliomurex chodmonensis n. sp.: I, holotype, MNHN.F.A81144 (Pacaud coll.), Chaumont-en-Vexin (Carrière Darcy), France, Lutetian; J, paratype, MNHN.F.A81145 (Pons coll.), Chaumont-en- Vexin (Carrière Darcy); K, L, Ponderia remyi n. sp.: K, holotype, MNHN.F.A90548 (Remy coll.), Fresville, Lutetian; L, paratype, MNHN.F.A90555, Fresville (Remy coll.); M, N, Flexopteron constantinense n. sp.: M, paratype, MNHN.F.A77761 (MNHN coll.), Carentan-les-Marais (Cauvin), France, Priabonian; N, holotype, MNHN.F.A77760 (MNHN coll.), Carentan-les-Marais (Cauvin). Scale bars: A-D, I-L, 10 mm; E-H, N, 5 mm; M, 3 mm. Credits: A, B, D-F, N, P. Loubry (MNHN/ CNRS); C, E. Robert (UCBL)." figureDoi="http://doi.org/10.5281/zenodo.13985831" httpUri="https://zenodo.org/record/13985831/files/figure.png" pageId="28" pageNumber="521">27D</figureCitation>
).
</materialsCitation>
</paragraph>
</subSubSection>
<subSubSection id="C33D6547FFFAE41AFC95F92BFCEFF8FA" pageId="28" pageNumber="521" type="etymology">
<paragraph id="8B9836CCFFFAE41AFC95F92BFCEFF8FA" blockId="28.[811,1462,1788,1865]" pageId="28" pageNumber="521">
<smallCapsWord id="8D7EA010FFFAE41AFC95F92BFC25F8A0" baselines="1806,1807" box="[813,932,1789,1812]" lowerCaseFontSize="7" mainFontSize="10" normCase="title" normString="Etymology" pageId="28" pageNumber="521">ETYMOLOGY</smallCapsWord>
. — Name formed by the prefix
<emphasis id="B953EADEFFFAE41AFB77F92AFB77F8A7" box="[1231,1270,1788,1812]" italics="true" pageId="28" pageNumber="521">sub-</emphasis>
and
<taxonomicName id="4C274D4FFFFAE41AFA9BF92BFA0FF8A7" baseAuthorityName="Deshayes" baseAuthorityYear="1835" box="[1315,1422,1789,1812]" class="Gastropoda" family="Muricidae" genus="Timbellus" kingdom="Animalia" order="Neogastropoda" pageId="28" pageNumber="521" phylum="Mollusca" rank="species" species="micropterus">
<emphasis id="B953EADEFFFAE41AFA9BF92BFA0FF8A7" box="[1315,1422,1789,1812]" italics="true" pageId="28" pageNumber="521">micropterus</emphasis>
</taxonomicName>
for this species older than Bartonian
<taxonomicName id="4C274D4FFFFAE41AFBC9F8C1FCEAF8FA" authority="(Deshayes, 1835)" baseAuthorityName="Deshayes" baseAuthorityYear="1835" class="Gastropoda" family="Muricidae" genus="Timbellus" kingdom="Animalia" order="Neogastropoda" pageId="28" pageNumber="521" phylum="Mollusca" rank="species" species="micropterus">
<emphasis id="B953EADEFFFAE41AFBC9F8C1FAC1F89D" box="[1137,1344,1815,1839]" italics="true" pageId="28" pageNumber="521">Timbellus micropterus</emphasis>
(
<bibRefCitation id="EFB64B3DFFFAE41AFAF7F8C1FCE2F8FA" author="DESHAYES G. - P." pageId="28" pageNumber="521" pagination="79 - 106" refId="ref43930" refString="DESHAYES G. - P. 1835. - Description des coquilles fossiles des environs de Paris. Tome 2. Livraisons 40 - 45, l'auteur, Bechet Jeune, Baudouin freres, Treuttel &amp; Wurtz, Paris: 499 - 780; Atlas 2, Levrault, Paris: pl. 79 - 106. https: // doi. org / 10.5962 / bhl. title. 52303" type="book chapter" year="1835">Deshayes, 1835</bibRefCitation>
)
</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection id="C33D6547FFFAE41AFC95F8BEFC41F85F" pageId="28" pageNumber="521" type="materials_examined">
<paragraph id="8B9836CCFFFAE41AFC95F8BEFBFDF829" blockId="28.[811,1456,1896,1946]" pageId="28" pageNumber="521">
<smallCapsWord id="8D7EA010FFFAE41AFC95F8BEFCDCF8CD" baselines="1913,1914" box="[813,861,1896,1919]" lowerCaseFontSize="7" mainFontSize="10" normCase="title" normString="Type" pageId="28" pageNumber="521">TYPE</smallCapsWord>
<smallCapsWord id="8D7EA010FFFAE41AFCDAF8BBFC3DF8CD" baselines="1914" box="[866,956,1901,1918]" lowerCaseFontSize="7" mainFontSize="10" normCase="lower" normString="horizon" pageId="28" pageNumber="521">HORIZON</smallCapsWord>
. — Calcaire de Fresville, middle Eocene (uppermost Lutetian), see
<bibRefCitation id="EFB64B3DFFFAE41AFC0EF854FBF7F829" author="BIGNOT G. &amp; HOMMERIL P. &amp; LARSONNEUR C." box="[950,1142,1922,1946]" pageId="28" pageNumber="521" pagination="405 - 416" refId="ref42941" refString="BIGNOT G., HOMMERIL P. &amp; LARSONNEUR C. 1968. - Le Lutetien au large du Cotentin, in Colloque sur l'Eocene (Paris, mai 1968). Memoires BRGM 58: 405 - 416." type="journal article" year="1968">
Bignot
<emphasis id="B953EADEFFFAE41AFC47F855FBAFF829" box="[1023,1070,1922,1946]" italics="true" pageId="28" pageNumber="521">et al.</emphasis>
(1968)
</bibRefCitation>
.
</paragraph>
<paragraph id="8B9836CCFFFAE41AFC95F86FFC41F85F" blockId="28.[813,1457,1977,2028]" pageId="28" pageNumber="521">
<smallCapsWord id="8D7EA010FFFAE41AFC95F86FFCE1F87C" baselines="1994,1995" box="[813,864,1977,2000]" lowerCaseFontSize="7" mainFontSize="10" normCase="title" normString="Type" pageId="28" pageNumber="521">TYPE</smallCapsWord>
<smallCapsWord id="8D7EA010FFFAE41AFCD1F868FC49F87C" baselines="1995" box="[873,968,1982,1999]" lowerCaseFontSize="7" mainFontSize="10" normCase="lower" normString="locality" pageId="28" pageNumber="521">LOCALITY</smallCapsWord>
. —
<collectingCountry id="F330765CFFFAE41AFC42F86FFBC0F863" box="[1018,1089,1977,2000]" name="France" pageId="28" pageNumber="521">France</collectingCountry>
,
<collectingRegion id="49E3F82EFFFAE41AFBF6F86FFB3EF862" box="[1102,1215,1977,2001]" country="France" name="Haute-Normandie" pageId="28" pageNumber="521">Normandy</collectingRegion>
, Manche, Fresville, see
<bibRefCitation id="EFB64B3DFFFAE41AFC97F805FC3DF85F" author="APGN &amp; ASSOCIATION PATRIMOINE GEOLOGIQUE DE NORMANDIE" box="[815,956,2003,2028]" pageId="28" pageNumber="521" refId="ref42597" refString="APGN (ASSOCIATION PATRIMOINE GEOLOGIQUE DE NORMANDIE) 2024. - Faluns cretaces et tertiaires de Fresville, BNO 0182. Available at: http: // www. apgn. fr / apgn / carte _ site / fic _ s 50 _ 75. pdf (last access on 03 / 2024)." type="book" year="2024">APGN (2024)</bibRefCitation>
.
</paragraph>
</subSubSection>
<subSubSection id="C33D6547FFFBE41BFF3CFF0EFD0CFE97" pageId="29" pageNumber="522" type="distribution">
<paragraph id="8B9836CCFFFBE41BFF3CFF0EFD0CFE97" blockId="29.[131,775,215,293]" pageId="29" pageNumber="522">
<smallCapsWord id="8D7EA010FFFBE41BFF3CFF0EFE91FF5D" baselines="233,234" box="[132,272,216,239]" lowerCaseFontSize="7" mainFontSize="10" normCase="title" normString="Distribution" pageId="29" pageNumber="522">DISTRIBUTION</smallCapsWord>
. —
<collectingRegion id="49E3F82EFFFBE41BFE81FF01FE22FF5C" box="[313,419,215,239]" country="France" name="Haute-Normandie" pageId="29" pageNumber="522">Normandy</collectingRegion>
(Calcaire de Fresville) middle Eocene (uppermost Lutetian) and?
<collectingRegion id="49E3F82EFFFBE41BFE26FF24FE50FEB9" box="[414,465,242,266]" country="France" name="Pays de la Loire" pageId="29" pageNumber="522">Loire</collectingRegion>
Basin (Sables du Bois-Gouët), lowermost Bartonian, see
<bibRefCitation id="EFB64B3DFFFBE41BFE39FEDAFD06FE96" author="MATHELIN J. - C. &amp; BIGNOT G." box="[385,647,268,293]" pageId="29" pageNumber="522" pagination="55 - 70" refId="ref46304" refString="MATHELIN J. - C. &amp; BIGNOT G. 1989. - Le falun de Foulangues et ses relations stratigraphique et paleogeographique avec le Biarritzien stratotypique, in 114 eme Congres national des Societes savantes. Geologie du Bassin Parisien. CTHS, Paris: 55 - 70." type="book chapter" year="1989">Mathelin &amp; Bignot (1989)</bibRefCitation>
.
</paragraph>
</subSubSection>
<subSubSection id="C33D6547FFFBE41BFF3CFE81FC86F858" pageId="29" pageNumber="522" type="description">
<paragraph id="8B9836CCFFFBE41BFF3CFE81FE94FEDC" blockId="29.[131,777,343,1039]" box="[132,277,343,369]" pageId="29" pageNumber="522">
<smallCapsWord id="8D7EA010FFFBE41BFF3CFE81FE94FEDC" baselines="362,362" box="[132,277,343,369]" lowerCaseFontSize="8" mainFontSize="11" normCase="title" normString="Description" pageId="29" pageNumber="522">DESCRIPTION</smallCapsWord>
</paragraph>
<paragraph id="8B9836CCFFFBE41BFF3CFEA1FDDFFBBC" blockId="29.[131,777,343,1039]" pageId="29" pageNumber="522">
Bulbous protoconch of one whorl (
<figureCitation id="131C2A49FFFBE41BFDB7FEA1FDF4FE22" box="[527,629,375,401]" captionStart="FIG" captionStartId="30.[132,143,1872,1889]" captionTargetBox="[147,1440,260,1687]" captionTargetId="figure-206@30.[700,1256,828,1396]" captionTargetPageId="30" captionText="FIG. 16. — Spiral cords and internal denticles of the outer lip in Timbellus submicropterus n. sp. (A-H, K) and T. micropterus (Deshayes, 1835) (I, J, L): A, B, holotype, MNHN.F.A24111 (MNHN coll.), Fresville, Manche, Normandy, France, middle Eocene (Lutetian); C, D, paratype, MNHN.F.A24126 (Munier-Chalmas coll.), Gourbesville, Manche, middle Eocene (Lutetian); E, F, paratype, UCBL-EM 33438 (Bourdot coll.), Fresville; G, H, MNHN.F.A90557 (Brébion coll.), Saint-Aignan- Grand-Lieu (Pierre-Aiguë), Loire-Atlantique, Loire Basin, France, middle Eocene (Bartonian); I, J, MNHN.F.A32522 (Faullummel coll.), Le Limon, Seine-et-Marne, Paris Basin, France,middle Eocene (Bartonian);K, paratype (protoconch), MNHN.F.A90556 (MNHN coll.), Fresville; L, MNHN.F.A90558 (Faullummel coll., protoconch), Chavençon (Carrière Tuyttens), Oise, Paris Basin, France, middle Eocene (Bartonian). Scale bars: A-J, 10 mm; K, L, 0.5 mm. Credits: A-D, I, J, P. Loubry (MNHN/CNRS); E, F, E. Robert (UCBL); G, H, K, L, L. Cazes (MNHN/CNRS)." figureDoi="http://doi.org/10.5281/zenodo.13985800" httpUri="https://zenodo.org/record/13985800/files/figure.png" pageId="29" pageNumber="522">Fig. 16K</figureCitation>
). Teleoconch up to
<quantity id="4CDF9B29FFFBE41BFF7BFE40FEA9FE02" box="[195,296,406,433]" metricMagnitude="-2" metricUnit="m" metricValue="2.36" pageId="29" pageNumber="522" unit="mm" value="23.6">23.6 mm</quantity>
in height, up to
<quantity id="4CDF9B29FFFBE41BFE67FE40FDC2FE02" box="[479,579,406,433]" metricMagnitude="-2" metricUnit="m" metricValue="1.16" pageId="29" pageNumber="522" unit="mm" value="11.6">11.6 mm</quantity>
in width, biconic in profile, composed of six whorls. Moderately high spire. Last whorl up to 70% of total length. Apical angle 59°. Spiral sculpture with coarse primary cords. First to third whorl: no cord. Fourth whorl: appearance of weak P1, s1 and P2. Fifth whorl: P1, s2 and P2 coarser. Sixth whorl: P1 to P6, s1; P6 not atrophied on convex part of whorl; ADP, MP and ABP on siphonal canal. From first whorl to second whorl: 6-7 protovarices. On third whorl: appearance of varices; three varices and one intervarix intercalated. From third to sixth whorl: no change. Higher relief of intervarices on P2. Varices rather thick. Appearance of short P1 spine on third whorl. Ovate aperture up to 27% of diameter and up to 71% of length of last whorl (including siphonal canal). Columellar lip smooth, narrow, poorly erect anteriorly, forming curved inductura at base. Parietal lip slightly adherent. Outer lip with developed denticles including ID, D1 to D6. D
<quantity id="4CDF9B29FFFBE41BFE0CFC63FE5FFC7C" box="[436,478,949,975]" metricMagnitude="-1" metricUnit="m" metricValue="1.524" pageId="29" pageNumber="522" unit="in" value="6.0">6 in</quantity>
some specimens. Outer lip not crenulate. Siphonal canal open, straight, up to 40% of apertural length. Pseudoumbilicus narrow.
</paragraph>
<paragraph id="8B9836CCFFFBE41BFF3CFBE2FEA2FBFE" blockId="29.[131,778,1076,2028]" box="[132,291,1076,1102]" pageId="29" pageNumber="522">
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</paragraph>
<paragraph id="8B9836CCFFFBE41BFF3CFB82FC86F858" blockId="29.[131,778,1076,2028]" pageId="29" pageNumber="522">
A specimen of this species (
<figureCitation id="131C2A49FFFBE41BFE12FB82FDA9FBDC" box="[426,552,1108,1135]" captionStart="FIG" captionStartId="30.[132,143,1872,1889]" captionTargetBox="[147,1440,260,1687]" captionTargetId="figure-206@30.[700,1256,828,1396]" captionTargetPageId="30" captionText="FIG. 16. — Spiral cords and internal denticles of the outer lip in Timbellus submicropterus n. sp. (A-H, K) and T. micropterus (Deshayes, 1835) (I, J, L): A, B, holotype, MNHN.F.A24111 (MNHN coll.), Fresville, Manche, Normandy, France, middle Eocene (Lutetian); C, D, paratype, MNHN.F.A24126 (Munier-Chalmas coll.), Gourbesville, Manche, middle Eocene (Lutetian); E, F, paratype, UCBL-EM 33438 (Bourdot coll.), Fresville; G, H, MNHN.F.A90557 (Brébion coll.), Saint-Aignan- Grand-Lieu (Pierre-Aiguë), Loire-Atlantique, Loire Basin, France, middle Eocene (Bartonian); I, J, MNHN.F.A32522 (Faullummel coll.), Le Limon, Seine-et-Marne, Paris Basin, France,middle Eocene (Bartonian);K, paratype (protoconch), MNHN.F.A90556 (MNHN coll.), Fresville; L, MNHN.F.A90558 (Faullummel coll., protoconch), Chavençon (Carrière Tuyttens), Oise, Paris Basin, France, middle Eocene (Bartonian). Scale bars: A-J, 10 mm; K, L, 0.5 mm. Credits: A-D, I, J, P. Loubry (MNHN/CNRS); E, F, E. Robert (UCBL); G, H, K, L, L. Cazes (MNHN/CNRS)." figureDoi="http://doi.org/10.5281/zenodo.13985800" httpUri="https://zenodo.org/record/13985800/files/figure.png" pageId="29" pageNumber="522">Figs 16E, F</figureCitation>
;
<figureCitation id="131C2A49FFFBE41BFD88FB82FDE3FBDD" box="[560,610,1108,1134]" captionStart="FIG" captionStartId="46.[132,143,1606,1623]" captionTargetBox="[132,1455,215,1546]" captionTargetId="figure-366@46.[699,1260,794,1359]" captionTargetPageId="46" captionText="FIG. 27. — New species of Timbellus De Gregorio, 1885, Pterynotus Swainson, 1833 (Pterymarchia Houart, 1995), Textiliomurex Merle, 2011, Ponderia Houart, 1986 and Flexopteron Shuto, 1969: A-C, Timbellus submicropterus n. sp.: A, holotype, MNHN.F.A24111 (MNHN coll.), Fresville, France, Lutetian; B, paratype, MNHN.F.A24126 (Munier-Chalmas coll.), Gourbesville, France, Lutetian; C, paratype, UCBL-EM 33438 (Bourdot coll.), Fresville; D, T. cf. submicropterus n. sp., MNHN.F.A90557 (Brébion coll.), Saint-Aignan-Grand-Lieu (Pierre-Aiguë), France, Bartonian; E, F, Pterynotus (Pterymarchia) gaasensis n. sp.: E, holotype, MNHN.F.A32492 (Merle coll.), Gaas (Espibos), France, Rupelian; F, paratype, MNHN.F.A90554 (Merle coll.), Gaas (Espibos); G, H, P. (P.) pelouatensis n. sp.: G, holotype, MNHN.F.J09945, Saucats (Peloua), France, Burdigalian; H, paratype, MNHN.F.J17632, Saucats (Peloua); I, J, Textiliomurex chodmonensis n. sp.: I, holotype, MNHN.F.A81144 (Pacaud coll.), Chaumont-en-Vexin (Carrière Darcy), France, Lutetian; J, paratype, MNHN.F.A81145 (Pons coll.), Chaumont-en- Vexin (Carrière Darcy); K, L, Ponderia remyi n. sp.: K, holotype, MNHN.F.A90548 (Remy coll.), Fresville, Lutetian; L, paratype, MNHN.F.A90555, Fresville (Remy coll.); M, N, Flexopteron constantinense n. sp.: M, paratype, MNHN.F.A77761 (MNHN coll.), Carentan-les-Marais (Cauvin), France, Priabonian; N, holotype, MNHN.F.A77760 (MNHN coll.), Carentan-les-Marais (Cauvin). Scale bars: A-D, I-L, 10 mm; E-H, N, 5 mm; M, 3 mm. Credits: A, B, D-F, N, P. Loubry (MNHN/ CNRS); C, E. Robert (UCBL)." figureDoi="http://doi.org/10.5281/zenodo.13985831" httpUri="https://zenodo.org/record/13985831/files/figure.png" pageId="29" pageNumber="522">27C</figureCitation>
) was identified as
<taxonomicName id="4C274D4FFFFBE41BFF1AFBA3FD39FB3D" authority="Lamarck, 1803" authorityName="Lamarck" authorityYear="1803" box="[162,696,1140,1166]" class="Gastropoda" family="Muricidae" genus="Murex" kingdom="Animalia" order="Neogastropoda" pageId="29" pageNumber="522" phylum="Mollusca" rank="species" species="tricarinatus" subGenus="Pterymurex">
<emphasis id="B953EADEFFFBE41BFF1AFBA3FF69FB3D" box="[162,232,1141,1166]" italics="true" pageId="29" pageNumber="522">Murex</emphasis>
(
<emphasis id="B953EADEFFFBE41BFF43FBA3FEF4FB3D" box="[251,373,1141,1166]" italics="true" pageId="29" pageNumber="522">Pterymurex</emphasis>
)
<emphasis id="B953EADEFFFBE41BFE3FFBA3FD85FB3D" box="[391,516,1141,1166]" italics="true" pageId="29" pageNumber="522">tricarinatus</emphasis>
<bibRefCitation id="EFB64B3DFFFBE41BFDB5FBA2FD39FB3D" author="LAMARCK J. - B." box="[525,696,1140,1166]" pageId="29" pageNumber="522" pagination="217 - 227" refId="ref45566" refString="LAMARCK J. - B. 1803. - Memoires sur les fossiles des environs de Paris (suite 1). Annales du Museum d'Histoire naturelle 2 (9): 217 - 227." type="journal article" year="1803">Lamarck, 1803</bibRefCitation>
</taxonomicName>
(synonym of
<taxonomicName id="4C274D4FFFFBE41BFF65FB42FDC1FB1D" baseAuthorityName="Lamarck" baseAuthorityYear="1803" box="[221,576,1172,1198]" class="Gastropoda" family="Muricidae" genus="Timbellus" kingdom="Animalia" order="Neogastropoda" pageId="29" pageNumber="522" phylum="Mollusca" rank="subSpecies" species="crenulatus" subSpecies="tricarinatus">
<emphasis id="B953EADEFFFBE41BFF65FB42FDC1FB1D" box="[221,576,1172,1198]" italics="true" pageId="29" pageNumber="522">Timbellus crenulatus tricarinatus</emphasis>
</taxonomicName>
) by
<bibRefCitation id="EFB64B3DFFFBE41BFDCCFB42FEB4FB7D" author="COSSMANN M. &amp; PISSARRO G." pageId="29" pageNumber="522" pagination="61 - 140" refId="ref43809" refString="COSSMANN M. &amp; PISSARRO G. 1901. - Faune eocenique du Cotentin. 2 eme article. Bulletin de la Societe Geologique de Normandie 20: 11 - 90. [Also published as Cossmann M. &amp; Pissarro G. 1901. - Faune eocenique du Cotentin (Mollusques), Tome 1, Fascicule 2, Imprimerie du Journal Le Havre, Le Havre, 61 - 140]." type="book chapter" year="1901">Cossmann &amp; Pissarro (1901)</bibRefCitation>
.
<taxonomicName id="4C274D4FFFFBE41BFEFFFB62FD36FB7D" baseAuthorityName="Lamarck" baseAuthorityYear="1803" box="[327,695,1204,1230]" class="Gastropoda" family="Muricidae" genus="Timbellus" kingdom="Animalia" order="Neogastropoda" pageId="29" pageNumber="522" phylum="Mollusca" rank="subSpecies" species="crenulatus" subSpecies="tricarinatus">
<emphasis id="B953EADEFFFBE41BFEFFFB62FD36FB7D" box="[327,695,1204,1230]" italics="true" pageId="29" pageNumber="522">Timbellus crenulatus tricarinatus</emphasis>
</taxonomicName>
differs greatly by apertural (with weak internal denticles within the outer lip), sculptural and shape characters, and belongs to another group species (see above
<figureCitation id="131C2A49FFFBE41BFDA5FAC5FDEAFA9D" box="[541,619,1299,1326]" captionStart="FIG" captionStartId="26.[132,143,1378,1395]" captionTargetBox="[132,1455,215,1334]" captionTargetId="figure-411@26.[698,1267,213,775]" captionTargetPageId="26" captionText="FIG. 14. — Spiral cords and internal denticles of the outer lip in Timbellus crenulatus tricarinatus (Lamarck, 1803) (A-C) and T. curvispina n. sp. (D-H): A, original figure of Lamarck (1816: pl. 418, fig. 5a), Thiverval-Grignon, Yvelines, Paris Basin, France, middle Eocene (Lutetian); B, C, neotype designated herein, MNHN.F.A90547 (MNHN coll.), Thiverval-Grignon (Falunière); D, paratype, MNHN.F.A73848 (Pons coll.), Fontenay-en-Vexin (Bois-du-But), Eure, Paris Basin, France,middle Eocene (Lutetian); E, F, holotype, MNHN.F.A90541 (MNHN coll.), Parnes, Oise, middle Eocene (Lutetian);G, H, paratype,MNHN.F.A90543 (Pons coll.), Fontenay-en-Vexin. Scale bars: B-H, 10 mm. Credits: P. Loubry." figureDoi="http://doi.org/10.5281/zenodo.13985794" httpUri="https://zenodo.org/record/13985794/files/figure.png" pageId="29" pageNumber="522">Fig. 14</figureCitation>
). In fact, this specimen illustrated by
<bibRefCitation id="EFB64B3DFFFBE41BFE3EFAE5FD47FAFE" author="COSSMANN M. &amp; PISSARRO G." box="[390,710,1331,1357]" pageId="29" pageNumber="522" pagination="61 - 140" refId="ref43809" refString="COSSMANN M. &amp; PISSARRO G. 1901. - Faune eocenique du Cotentin. 2 eme article. Bulletin de la Societe Geologique de Normandie 20: 11 - 90. [Also published as Cossmann M. &amp; Pissarro G. 1901. - Faune eocenique du Cotentin (Mollusques), Tome 1, Fascicule 2, Imprimerie du Journal Le Havre, Le Havre, 61 - 140]." type="book chapter" year="1901">Cossmann &amp; Pissarro (1901)</bibRefCitation>
more closely resembles
<taxonomicName id="4C274D4FFFFBE41BFEF5FA82FE66FADE" baseAuthorityName="Deshayes" baseAuthorityYear="1835" box="[333,487,1364,1389]" class="Gastropoda" family="Muricidae" genus="Timbellus" kingdom="Animalia" order="Neogastropoda" pageId="29" pageNumber="522" phylum="Mollusca" rank="species" species="micropterus">
<emphasis id="B953EADEFFFBE41BFEF5FA82FE66FADE" box="[333,487,1364,1389]" italics="true" pageId="29" pageNumber="522">T. micropterus</emphasis>
</taxonomicName>
from the Bartonian Paris Basin.
<bibRefCitation id="EFB64B3DFFFBE41BFF75FAA5FE15FA3E" author="MERLE D. &amp; GARRIGUES B. &amp; POINTIER J. - P." box="[205,404,1395,1421]" pageId="29" pageNumber="522" refId="ref47234" refString="MERLE D., GARRIGUES B. &amp; POINTIER J. - P. 2011. - Fossil &amp; Recent Muricidae of the World. Part Muricinae. ConchBooks, Hackenheim, 652 p." type="book" year="2011">
<taxonomicName id="4C274D4FFFFBE41BFF75FAA5FE8CFA3E" authorityName="Merle &amp; Pacaud &amp; Ledon &amp; Goret" authorityYear="2024" box="[205,269,1395,1421]" class="Gastropoda" family="Muricidae" kingdom="Animalia" order="Neogastropoda" pageId="29" pageNumber="543" phylum="Mollusca" rank="subFamily" status="subfam. nov." subFamily="Merle">Merle</taxonomicName>
<emphasis id="B953EADEFFFBE41BFEAAFAA2FEC4FA3E" box="[274,325,1395,1421]" italics="true" pageId="29" pageNumber="522">et al.</emphasis>
(2011)
</bibRefCitation>
attributed another Norman specimen (pl. 102, fig. 4a, b) to
<taxonomicName id="4C274D4FFFFBE41BFE7EFA42FDE0FA1E" baseAuthorityName="Deshayes" baseAuthorityYear="1835" box="[454,609,1428,1453]" class="Gastropoda" family="Muricidae" genus="Timbellus" kingdom="Animalia" order="Neogastropoda" pageId="29" pageNumber="522" phylum="Mollusca" rank="species" species="micropterus">
<emphasis id="B953EADEFFFBE41BFE7EFA42FDE0FA1E" box="[454,609,1428,1453]" italics="true" pageId="29" pageNumber="522">T. micropterus</emphasis>
</taxonomicName>
(
<figureCitation id="131C2A49FFFBE41BFDCAFA45FC83FA1D" box="[626,770,1427,1454]" captionStart="FIG" captionStartId="30.[132,143,1872,1889]" captionTargetBox="[147,1440,260,1687]" captionTargetId="figure-206@30.[700,1256,828,1396]" captionTargetPageId="30" captionText="FIG. 16. — Spiral cords and internal denticles of the outer lip in Timbellus submicropterus n. sp. (A-H, K) and T. micropterus (Deshayes, 1835) (I, J, L): A, B, holotype, MNHN.F.A24111 (MNHN coll.), Fresville, Manche, Normandy, France, middle Eocene (Lutetian); C, D, paratype, MNHN.F.A24126 (Munier-Chalmas coll.), Gourbesville, Manche, middle Eocene (Lutetian); E, F, paratype, UCBL-EM 33438 (Bourdot coll.), Fresville; G, H, MNHN.F.A90557 (Brébion coll.), Saint-Aignan- Grand-Lieu (Pierre-Aiguë), Loire-Atlantique, Loire Basin, France, middle Eocene (Bartonian); I, J, MNHN.F.A32522 (Faullummel coll.), Le Limon, Seine-et-Marne, Paris Basin, France,middle Eocene (Bartonian);K, paratype (protoconch), MNHN.F.A90556 (MNHN coll.), Fresville; L, MNHN.F.A90558 (Faullummel coll., protoconch), Chavençon (Carrière Tuyttens), Oise, Paris Basin, France, middle Eocene (Bartonian). Scale bars: A-J, 10 mm; K, L, 0.5 mm. Credits: A-D, I, J, P. Loubry (MNHN/CNRS); E, F, E. Robert (UCBL); G, H, K, L, L. Cazes (MNHN/CNRS)." figureDoi="http://doi.org/10.5281/zenodo.13985800" httpUri="https://zenodo.org/record/13985800/files/figure.png" pageId="29" pageNumber="522">Figs 16C, D</figureCitation>
;
<figureCitation id="131C2A49FFFBE41BFF3EFA65FF37FA7E" box="[134,182,1459,1485]" captionStart="FIG" captionStartId="46.[132,143,1606,1623]" captionTargetBox="[132,1455,215,1546]" captionTargetId="figure-366@46.[699,1260,794,1359]" captionTargetPageId="46" captionText="FIG. 27. — New species of Timbellus De Gregorio, 1885, Pterynotus Swainson, 1833 (Pterymarchia Houart, 1995), Textiliomurex Merle, 2011, Ponderia Houart, 1986 and Flexopteron Shuto, 1969: A-C, Timbellus submicropterus n. sp.: A, holotype, MNHN.F.A24111 (MNHN coll.), Fresville, France, Lutetian; B, paratype, MNHN.F.A24126 (Munier-Chalmas coll.), Gourbesville, France, Lutetian; C, paratype, UCBL-EM 33438 (Bourdot coll.), Fresville; D, T. cf. submicropterus n. sp., MNHN.F.A90557 (Brébion coll.), Saint-Aignan-Grand-Lieu (Pierre-Aiguë), France, Bartonian; E, F, Pterynotus (Pterymarchia) gaasensis n. sp.: E, holotype, MNHN.F.A32492 (Merle coll.), Gaas (Espibos), France, Rupelian; F, paratype, MNHN.F.A90554 (Merle coll.), Gaas (Espibos); G, H, P. (P.) pelouatensis n. sp.: G, holotype, MNHN.F.J09945, Saucats (Peloua), France, Burdigalian; H, paratype, MNHN.F.J17632, Saucats (Peloua); I, J, Textiliomurex chodmonensis n. sp.: I, holotype, MNHN.F.A81144 (Pacaud coll.), Chaumont-en-Vexin (Carrière Darcy), France, Lutetian; J, paratype, MNHN.F.A81145 (Pons coll.), Chaumont-en- Vexin (Carrière Darcy); K, L, Ponderia remyi n. sp.: K, holotype, MNHN.F.A90548 (Remy coll.), Fresville, Lutetian; L, paratype, MNHN.F.A90555, Fresville (Remy coll.); M, N, Flexopteron constantinense n. sp.: M, paratype, MNHN.F.A77761 (MNHN coll.), Carentan-les-Marais (Cauvin), France, Priabonian; N, holotype, MNHN.F.A77760 (MNHN coll.), Carentan-les-Marais (Cauvin). Scale bars: A-D, I-L, 10 mm; E-H, N, 5 mm; M, 3 mm. Credits: A, B, D-F, N, P. Loubry (MNHN/ CNRS); C, E. Robert (UCBL)." figureDoi="http://doi.org/10.5281/zenodo.13985831" httpUri="https://zenodo.org/record/13985831/files/figure.png" pageId="29" pageNumber="522">27B</figureCitation>
). It is true that these
<specimenCount id="9D21FD45FFFBE41BFE14FA65FDD1FA7E" box="[428,592,1459,1485]" count="2" pageId="29" pageNumber="522" type="generic">two specimens</specimenCount>
are superficially similar to
<taxonomicName id="4C274D4FFFFBE41BFF45FA02FE19FA5E" baseAuthorityName="Deshayes" baseAuthorityYear="1835" box="[253,408,1492,1517]" class="Gastropoda" family="Muricidae" genus="Timbellus" kingdom="Animalia" order="Neogastropoda" pageId="29" pageNumber="522" phylum="Mollusca" rank="species" species="micropterus">
<emphasis id="B953EADEFFFBE41BFF45FA02FE19FA5E" box="[253,408,1492,1517]" italics="true" pageId="29" pageNumber="522">T. micropterus</emphasis>
</taxonomicName>
, but further comparison reveals some differences, such as only one intervarice (instead two in typical
<taxonomicName id="4C274D4FFFFBE41BFE89F9C5FE4CF99F" baseAuthorityName="Deshayes" baseAuthorityYear="1835" box="[305,461,1555,1580]" class="Gastropoda" family="Muricidae" genus="Timbellus" kingdom="Animalia" order="Neogastropoda" pageId="29" pageNumber="522" phylum="Mollusca" rank="species" species="micropterus">
<emphasis id="B953EADEFFFBE41BFE89F9C5FE4CF99F" box="[305,461,1555,1580]" italics="true" pageId="29" pageNumber="522">T. micropterus</emphasis>
</taxonomicName>
) and finer secondary cords on the last whorl. The bivariate plot of total length against width (
<figureCitation id="131C2A49FFFBE41BFF60F984FEA8F9DF" box="[216,297,1618,1644]" captionStart="FIG" captionStartId="31.[131,142,891,908]" captionTargetBox="[269,1278,178,858]" captionTargetId="figure-686@31.[347,1001,217,720]" captionText="FIG. 17. — Bivariate plot of total length against width comparing T. micropterus (Deshayes, 1835) from the lower Bartonian of the Paris Basin (50 complete specimens) and T. submicropterus n. sp. from the uppper Lutetian of Normandy (five complete specimens)." figureDoi="http://doi.org/10.5281/zenodo.13988488" httpUri="https://zenodo.org/record/13988488/files/figure.png" pageId="29" pageNumber="522">Fig. 17</figureCitation>
) does not suggest strong differences
<collectingRegion id="49E3F82EFFFBE41BFD69F985FF51F93F" country="France" name="Haute-Normandie" pageId="29" pageNumber="522">Normandy</collectingRegion>
and Paris Basin populations. Finally, it is tempting to conclude that the observed teleoconch differences in
<taxonomicName id="4C274D4FFFFBE41BFD49F945FEA2F97F" authorityName="Merle &amp; Pacaud &amp; Ledon &amp; Goret" authorityYear="2024" class="Gastropoda" family="Muricidae" genus="Timbellus" kingdom="Animalia" order="Neogastropoda" pageId="29" pageNumber="522" phylum="Mollusca" rank="species" species="submicropterus" status="n. sp.">
<emphasis id="B953EADEFFFBE41BFD49F945FEA2F97F" italics="true" pageId="29" pageNumber="522">T. submicropterus</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A26057A5FFFBE41BFE94F965FEE8F97E" box="[300,361,1715,1741]" pageId="29" pageNumber="522" rank="species">n. sp.</taxonomicNameLabel>
fall within the intraspecific variation of
<taxonomicName id="4C274D4FFFFBE41BFF1BF905FEBFF95F" baseAuthorityName="Deshayes" baseAuthorityYear="1835" box="[163,318,1747,1772]" class="Gastropoda" family="Muricidae" genus="Timbellus" kingdom="Animalia" order="Neogastropoda" pageId="29" pageNumber="522" phylum="Mollusca" rank="species" species="micropterus">
<emphasis id="B953EADEFFFBE41BFF1BF905FEBFF95F" box="[163,318,1747,1772]" italics="true" pageId="29" pageNumber="522">T. micropterus</emphasis>
</taxonomicName>
. However,
<taxonomicName id="4C274D4FFFFBE41BFE7AF905FD01F95F" authorityName="Merle &amp; Pacaud &amp; Ledon &amp; Goret" authorityYear="2024" box="[450,640,1746,1772]" class="Gastropoda" family="Muricidae" genus="Timbellus" kingdom="Animalia" order="Neogastropoda" pageId="29" pageNumber="522" phylum="Mollusca" rank="species" species="submicropterus" status="n. sp.">
<emphasis id="B953EADEFFFBE41BFE7AF905FD01F95F" box="[450,640,1746,1772]" italics="true" pageId="29" pageNumber="522">T. submicropterus</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A26057A5FFFBE41BFD32F904FD49F95F" box="[650,712,1746,1772]" pageId="29" pageNumber="522" rank="species">n. sp.</taxonomicNameLabel>
has a paucispiral protoconch, whereas
<taxonomicName id="4C274D4FFFFBE41BFE4AF925FD0DF8BF" baseAuthorityName="Deshayes" baseAuthorityYear="1835" box="[498,652,1779,1804]" class="Gastropoda" family="Muricidae" genus="Timbellus" kingdom="Animalia" order="Neogastropoda" pageId="29" pageNumber="522" phylum="Mollusca" rank="species" species="micropterus">
<emphasis id="B953EADEFFFBE41BFE4AF925FD0DF8BF" box="[498,652,1779,1804]" italics="true" pageId="29" pageNumber="522">T. micropterus</emphasis>
</taxonomicName>
has a multispiral (
<figureCitation id="131C2A49FFFBE41BFF59F8C4FEE5F89F" box="[225,356,1810,1836]" captionStart="FIG" captionStartId="30.[132,143,1872,1889]" captionTargetBox="[147,1440,260,1687]" captionTargetId="figure-206@30.[700,1256,828,1396]" captionTargetPageId="30" captionText="FIG. 16. — Spiral cords and internal denticles of the outer lip in Timbellus submicropterus n. sp. (A-H, K) and T. micropterus (Deshayes, 1835) (I, J, L): A, B, holotype, MNHN.F.A24111 (MNHN coll.), Fresville, Manche, Normandy, France, middle Eocene (Lutetian); C, D, paratype, MNHN.F.A24126 (Munier-Chalmas coll.), Gourbesville, Manche, middle Eocene (Lutetian); E, F, paratype, UCBL-EM 33438 (Bourdot coll.), Fresville; G, H, MNHN.F.A90557 (Brébion coll.), Saint-Aignan- Grand-Lieu (Pierre-Aiguë), Loire-Atlantique, Loire Basin, France, middle Eocene (Bartonian); I, J, MNHN.F.A32522 (Faullummel coll.), Le Limon, Seine-et-Marne, Paris Basin, France,middle Eocene (Bartonian);K, paratype (protoconch), MNHN.F.A90556 (MNHN coll.), Fresville; L, MNHN.F.A90558 (Faullummel coll., protoconch), Chavençon (Carrière Tuyttens), Oise, Paris Basin, France, middle Eocene (Bartonian). Scale bars: A-J, 10 mm; K, L, 0.5 mm. Credits: A-D, I, J, P. Loubry (MNHN/CNRS); E, F, E. Robert (UCBL); G, H, K, L, L. Cazes (MNHN/CNRS)." figureDoi="http://doi.org/10.5281/zenodo.13985800" httpUri="https://zenodo.org/record/13985800/files/figure.png" pageId="29" pageNumber="522">Fig. 16K, L</figureCitation>
), suggesting two separate species with different modes of larval development. In the Bartonian from Loire-Atlantique, there are specimens showing coarse spiral cords like those of typical
<taxonomicName id="4C274D4FFFFBE41BFE42F8A4FD3BF838" authorityName="Merle &amp; Pacaud &amp; Ledon &amp; Goret" authorityYear="2024" box="[506,698,1905,1931]" class="Gastropoda" family="Muricidae" genus="Timbellus" kingdom="Animalia" order="Neogastropoda" pageId="29" pageNumber="522" phylum="Mollusca" rank="species" species="submicropterus" status="n. sp.">
<emphasis id="B953EADEFFFBE41BFE42F8A4FD3BF838" box="[506,698,1905,1931]" italics="true" pageId="29" pageNumber="522">T. submicropterus</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A26057A5FFFBE41BFD7BF8A4FC85F83F" box="[707,772,1906,1932]" pageId="29" pageNumber="522" rank="species">n. sp.</taxonomicNameLabel>
, but they bear a more pronounced shoulder (
<figureCitation id="131C2A49FFFBE41BFDD6F847FD79F818" box="[622,760,1937,1963]" captionStart="FIG" captionStartId="30.[132,143,1872,1889]" captionTargetBox="[147,1440,260,1687]" captionTargetId="figure-206@30.[700,1256,828,1396]" captionTargetPageId="30" captionText="FIG. 16. — Spiral cords and internal denticles of the outer lip in Timbellus submicropterus n. sp. (A-H, K) and T. micropterus (Deshayes, 1835) (I, J, L): A, B, holotype, MNHN.F.A24111 (MNHN coll.), Fresville, Manche, Normandy, France, middle Eocene (Lutetian); C, D, paratype, MNHN.F.A24126 (Munier-Chalmas coll.), Gourbesville, Manche, middle Eocene (Lutetian); E, F, paratype, UCBL-EM 33438 (Bourdot coll.), Fresville; G, H, MNHN.F.A90557 (Brébion coll.), Saint-Aignan- Grand-Lieu (Pierre-Aiguë), Loire-Atlantique, Loire Basin, France, middle Eocene (Bartonian); I, J, MNHN.F.A32522 (Faullummel coll.), Le Limon, Seine-et-Marne, Paris Basin, France,middle Eocene (Bartonian);K, paratype (protoconch), MNHN.F.A90556 (MNHN coll.), Fresville; L, MNHN.F.A90558 (Faullummel coll., protoconch), Chavençon (Carrière Tuyttens), Oise, Paris Basin, France, middle Eocene (Bartonian). Scale bars: A-J, 10 mm; K, L, 0.5 mm. Credits: A-D, I, J, P. Loubry (MNHN/CNRS); E, F, E. Robert (UCBL); G, H, K, L, L. Cazes (MNHN/CNRS)." figureDoi="http://doi.org/10.5281/zenodo.13985800" httpUri="https://zenodo.org/record/13985800/files/figure.png" pageId="29" pageNumber="522">Fig. 16G, H</figureCitation>
). Their protoconch is not preserved. We therefore attribute these specimens to
<taxonomicName id="4C274D4FFFFBE41BFEEDF804FD90F858" authorityName="Merle &amp; Pacaud &amp; Ledon &amp; Goret" authorityYear="2024" box="[341,529,2001,2027]" class="Gastropoda" family="Muricidae" genus="Timbellus" kingdom="Animalia" order="Neogastropoda" pageId="29" pageNumber="522" phylum="Mollusca" rank="species" species="submicropterus" status="n. sp.">
<emphasis id="B953EADEFFFBE41BFEEDF804FD90F858" box="[341,529,2001,2027]" italics="true" pageId="29" pageNumber="522">T. submicropterus</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A26057A5FFFBE41BFDA0F804FDD5F85F" box="[536,596,2002,2028]" pageId="29" pageNumber="522" rank="species">n. sp.</taxonomicNameLabel>
with hesitation.
</paragraph>
</subSubSection>
</treatment>
</document>