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<document id="4645A45250AFD2C5A282050C5536227A" ID-DOI="10.5252/geodiversitas2019v41a5" ID-GBIF-Dataset="1dd366f3-9e6d-4dcf-a082-408a0ea91bc4" ID-ISSN="1638-9395" ID-Zenodo-Dep="3699932" ID-ZooBank="urn:lsid:zoobank.org:pub:33E8EF32-4BE4-4992-9917-8611CA0AD579" IM.bibliography_approvedBy="felipe" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" IM.treatmentCitations_approvedBy="felipe" IM.treatments_approvedBy="felipe" checkinTime="1583514705026" checkinUser="valdenar" docAuthor="Sanders, Malcolm T., Merle, Didier &amp; Puillandre, Nicolas" docDate="2019" docId="03D387DF1938FFEDFF3FFF2AFD28FCE3" docLanguage="en" docName="GEODIVERSITAS.41.5.247-265.pdf.imf" docOrigin="Geodiversitas 41 (5)" docStyle="DocumentStyle:F830B10FF475E64C1F1601E3B32DDC00.4:Geodiversitas.2018-.journal_article" docStyleId="F830B10FF475E64C1F1601E3B32DDC00" docStyleName="Geodiversitas.2018-.journal_article" docStyleVersion="4" docTitle="Aquitanobursa tuberosa Sanders &amp; Merle &amp; Puillandre 2019, n. comb" docType="treatment" docVersion="5" lastPageNumber="261" masterDocId="FFEAFFA71936FFFDFF96FFF2FFC1FA0D" masterDocTitle="A review of fossil Bursidae and their use for phylogeny calibration" masterLastPageNumber="265" masterPageNumber="247" pageNumber="259" updateTime="1698725915702" updateUser="plazi" zenodo-license-document="CC0-1.0" zenodo-license-figures="CC0-1.0">
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<mods:title id="54C35AD1E9BD69A9273752B7157AC2FB">A review of fossil Bursidae and their use for phylogeny calibration</mods:title>
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<mods:namePart id="05E99B89C108B03DA0133A79D5702E70">Sanders, Malcolm T.</mods:namePart>
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<heading id="D08D81A51938FFF3FF3FFF2AFD1EFAFE" box="[169,735,216,244]" pageId="14" pageNumber="259">
<taxonomicName id="4C7A4D4A1938FFF3FF3FFF2AFDBBFAFE" ID-CoL="7Q6LZ" authority="(Grateloup, 1833)" authorityName="Sanders &amp; Merle &amp; Puillandre" authorityYear="2019" baseAuthorityName="Grateloup" baseAuthorityYear="1833" box="[169,634,216,243]" class="Gastropoda" family="Bursidae" genus="Aquitanobursa" higherTaxonomySource="GBIF" kingdom="Animalia" order="Littorinimorpha" pageId="14" pageNumber="259" phylum="Mollusca" rank="species" species="tuberosa" status="comb. nov.">
<emphasis id="B90EEADB1938FFF3FF3FFF2AFE6CFAFE" bold="true" box="[169,429,216,243]" italics="true" pageId="14" pageNumber="259">Aquitanobursa tuberosa</emphasis>
(
<bibRefCitation id="EFEB4B381938FFF3FE2DFF2BFDB3FAFE" author="GRATELOUP J. P. S. DE" box="[443,626,217,243]" pageId="14" pageNumber="259" pagination="159 - 164" refId="ref11170" refString="GRATELOUP J. P. S. DE 1833. - Tableau des coquilles fossiles qu'on rencontre dans les terrains calcaire tertiaires (faluns) des environs de Dax, departement des Landes. Actes de la Societe linneenne de Bordeaux 6: 159 - 164. https: // biodiversitylibrary. org / page / 32986064" type="journal article" year="1833">Grateloup, 1833</bibRefCitation>
)
</taxonomicName>
<taxonomicNameLabel id="A23D57A01938FFF3FD17FF28FD1EFAFE" box="[641,735,217,244]" pageId="14" pageNumber="259" rank="species">n. comb</taxonomicNameLabel>
</heading>
. (
<figureCitation id="13412A4C1938FFF3FE0AFF0BFE2EFB1E" box="[412,495,249,276]" captionStart="FIG" captionStartId="12.[132,143,1964,1982]" captionTargetBox="[147,1440,215,1878]" captionTargetId="figure@12.[701,1257,768,1324]" captionTargetPageId="12" captionText="FIG. 7. — Aquitanobursa n. gen. of Aquitaine Basin: A, Aquitanobursa grateloupi (dOrbigny, 1852) n. comb., MNHN.F.A27211, lectotype from the Aquitanian of Saint-Paul-les-Dax, France; B, Aquitanobursa inaequicrenata (Cossmann &amp; Peyrot,1924) n. comb., MNHN.F.A70284, Burdigalian of le Peloua, France; Faullummel coll.; C, Aquitanobursa tuberosa (Grateloup, 1833) n. comb., MNHN.F.A70285, Burdigalian of le Peloua, Staadt coll." figureDoi="http://doi.org/10.5281/zenodo.3699946" httpUri="https://zenodo.org/record/3699946/files/figure.png" pageId="14" pageNumber="259">Fig. 7C</figureCitation>
)
<treatmentCitationGroup id="AB6A11E71938FFF3FF12FEC0FE46FBD3" pageId="14" pageNumber="259">
<treatmentCitation id="0ADB10D81938FFF3FF12FEC0FE35FB47" author="GRATELOUP J. P. S. DE" box="[132,500,306,330]" page="92" pageId="14" pageNumber="259" year="1833">
<taxonomicName id="4C7A4D4A1938FFF3FF12FEC0FE35FB47" authority="Grateloup, 1833: 92" authorityName="Grateloup" authorityPageNumber="92" authorityYear="1833" box="[132,500,306,330]" class="Gastropoda" family="Ranellidae" genus="Ranella" kingdom="Animalia" order="Littorinimorpha" pageId="14" pageNumber="259" phylum="Mollusca" rank="species" species="tuberosa">
<emphasis id="B90EEADB1938FFF3FF12FEC0FEE1FB47" box="[132,288,306,330]" italics="true" pageId="14" pageNumber="259">Ranella tuberosa</emphasis>
<treatmentCitation id="0ADB10D81938FFF3FEB1FEC0FE35FB47" author="GRATELOUP J. P. S. DE" box="[295,500,306,330]" page="" pageId="14" pageNumber="259" year="1833">
<bibRefCitation id="EFEB4B381938FFF3FEB1FEC0FE35FB47" author="GRATELOUP J. P. S. DE" box="[295,500,306,330]" pageId="14" pageNumber="259" pagination="159 - 164" refId="ref11170" refString="GRATELOUP J. P. S. DE 1833. - Tableau des coquilles fossiles qu'on rencontre dans les terrains calcaire tertiaires (faluns) des environs de Dax, departement des Landes. Actes de la Societe linneenne de Bordeaux 6: 159 - 164. https: // biodiversitylibrary. org / page / 32986064" type="journal article" year="1833">Grateloup, 1833: 92</bibRefCitation>
</treatmentCitation>
</taxonomicName>
</treatmentCitation>
, no. 420. —
<treatmentCitation id="0ADB10D81938FFF3FDEDFEC0FF6DFB69" author="BELLARDI L." page="236" pageId="14" pageNumber="259" year="1873">
<bibRefCitation id="EFEB4B381938FFF3FDEDFEC0FCC0FB47" author="BELLARDI L." box="[635,769,306,330]" pageId="14" pageNumber="259" pagination="33 - 294" refId="ref10139" refString="BELLARDI L. 1873. - I molluschi dei terreni Terziarii del Piemonte e della Liguria. Part 1: Cephalopoda. Pteropoda. Heteropoda. Gasteropoda (Muricidae e Tritonidae). Memorie della Reale Accademia delle Scienze di Torino ser. 2, 27: 33 - 294. https: // biodiversitylibrary. org / page / 12135674" type="journal article" year="1873">Bellardi 1873</bibRefCitation>
: 236
</treatmentCitation>
, pl. 15, fig. 7.
<treatmentCitation id="0ADB10D81938FFF3FF12FE8EFDD1FB99" box="[132,528,380,404]" pageId="14" pageNumber="259">
<taxonomicName id="4C7A4D4A1938FFF3FF12FE8EFE2BFB99" ID-CoL="7PFK2" authority="d'Orbigny, 1852" authorityName="d'Orbigny" authorityYear="1852" box="[132,490,380,404]" class="Gastropoda" family="Ranellidae" genus="Ranella" kingdom="Animalia" order="Littorinimorpha" pageId="14" pageNumber="250" phylum="Mollusca" rank="genus">
<emphasis id="B90EEADB1938FFF3FF12FE8EFEFFFB99" box="[132,318,380,404]" italics="true" pageId="14" pageNumber="259">Ranella subtuberosa</emphasis>
dOrbigny, 1852
</taxonomicName>
: 76
</treatmentCitation>
, no. 1405.
<emphasis id="B90EEADB1938FFF3FF12FE59FE85FBCE" box="[132,324,427,451]" italics="true" pageId="14" pageNumber="259">Apollon pelouatensis</emphasis>
<treatmentCitation id="0ADB10D81938FFF3FEDAFE5EFD9CFBC9" author="COSSMANN M. &amp; PEYROT A." box="[332,605,428,452]" page="" pageId="14" pageNumber="259" year="1924">
<bibRefCitation id="EFEB4B381938FFF3FEDAFE5EFD9CFBC9" author="COSSMANN M. &amp; PEYROT A." box="[332,605,428,452]" pageId="14" pageNumber="259" pagination="193 - 318" refId="ref10623" refString="COSSMANN M. &amp; PEYROT A. 1924. - Conchologie neogenique de l'Aquitaine (Suite). Actes de la Societe linneenne de Bordeaux 75 (3): 193 - 318. https: // biodiversitylibrary. org / page / 32207388" type="journal article" year="1924">Cossmann &amp; Peyrot, 1924</bibRefCitation>
</treatmentCitation>
: no. 842, pl. 15, figs 38-39, pl. 17, figs 1-2.
</treatmentCitationGroup>
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</paragraph>
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<subSubSection id="C36065421938FFF3FF12FE04FBCDFE9C" pageId="14" pageNumber="259" type="reference_group">
<paragraph id="8BC536C91938FFF3FF12FE04FD6EFEBD" blockId="14.[132,783,216,1108]" lastBlockId="14.[132,776,1174,2030]" pageId="14" pageNumber="259">
<taxonomicName id="4C7A4D4A1938FFF3FF12FE04FEFEF803" authority="" box="[132,319,502,526]" class="Gastropoda" family="Bursidae" genus="Bursa" kingdom="Animalia" order="Littorinimorpha" pageId="14" pageNumber="259" phylum="Mollusca" rank="subGenus" subGenus="Bufonariella">
<emphasis id="B90EEADB1938FFF3FF12FE04FF78F800" box="[132,185,502,525]" italics="true" pageId="14" pageNumber="259">Bursa</emphasis>
(
<emphasis id="B90EEADB1938FFF3FF53FE04FEF6F803" box="[197,311,502,526]" italics="true" pageId="14" pageNumber="259">Bufonariella</emphasis>
)
</taxonomicName>
<emphasis id="B90EEADB1938FFF3FED2FE04FE6EF803" box="[324,431,502,526]" italics="true" pageId="14" pageNumber="259">pelouatensis</emphasis>
<treatmentCitation id="0ADB10D81938FFF3FE53FE04FD95F800" author="BEU A. G." box="[453,596,501,526]" page="258" pageId="14" pageNumber="259" year="1981">
<bibRefCitation id="EFEB4B381938FFF3FE53FE04FD95F800" author="BEU A. G." box="[453,596,501,526]" pageId="14" pageNumber="259" pagination="248 - 324" refId="ref10200" refString="BEU A. G. 1981. - Australian gastropods of the family Bursidae. Part 1. The families of Tonnacea, the genera of Bursidae, and revision of species previously assigned to Tutufa Jousseaume, 1881. Records of the Australian Museum 33: 248 - 324. https: // doi. org / 10.3853 / j. 0067 - 1975.33.1980.200" type="journal article" year="1981">Beu 1981: 258</bibRefCitation>
</treatmentCitation>
. —
<treatmentCitation id="0ADB10D81938FFF3FDECFE04FD6EFEBD" author="SCHMELZ G. W." page="106" pageId="14" pageNumber="259" year="1997">
<bibRefCitation id="EFEB4B381938FFF3FDECFE04FCC0F803" author="SCHMELZ G. W." box="[634,769,502,526]" pageId="14" pageNumber="259" pagination="105 - 108" refId="ref12815" refString="SCHMELZ G. W. 1997. - Notes on the fauna of the Chipola Formation - XLI. A reexamination of Bursa (Mollusca: Gastropoda) from the Chipola Formation, with the description of a new species. Tulane Studies in Geology and Paleontology 30: 105 - 108." type="journal article" year="1997">Schmelz 1997</bibRefCitation>
:
<bibRefCitation id="EFEB4B381938FFF3FF12FDE2FD6EFEBD" author="BEU A. G." pageId="14" pageNumber="259" pagination="1 - 186" refId="ref10478" refString="BEU A. G. 2005. - Neogene fossil tonnoidean gastropods of Indonesia. Scripta Geologica 130: 1 - 186." type="journal article" year="2005">
106, pl. 1, fig. 2.
<taxonomicName id="4C7A4D4A1938FFF3FF12FDB2FECDF855" box="[132,268,576,600]" class="Gastropoda" family="Bursidae" genus="Bursa" kingdom="Animalia" order="Littorinimorpha" pageId="14" pageNumber="259" phylum="Mollusca" rank="species" species="tuberosa">
<emphasis id="B90EEADB1938FFF3FF12FDB2FECDF855" box="[132,268,576,600]" italics="true" pageId="14" pageNumber="259">Bursa tuberosa</emphasis>
</taxonomicName>
<treatmentCitation id="0ADB10D81938FFF3FEB5FDB2FDC5F855" author="LOZOUET P. &amp; LESPORT J. - F. &amp; RENARD P." box="[291,516,576,600]" page="45" pageId="14" pageNumber="259" year="2005">
<bibRefCitation id="EFEB4B381938FFF3FEB5FDB2FDC5F855" author="LOZOUET P. &amp; LESPORT J. - F. &amp; RENARD P." box="[291,516,576,600]" pageId="14" pageNumber="259" pagination="1 - 189" refId="ref11962" refString="LOZOUET P., LESPORT J. - F. &amp; RENARD P. 2001. - Revision des gastropoda (Mollusca) du Stratotype de l'Aquitanien. Cossmanniana (h. s. 3): 1 - 189." type="book chapter" year="2001">
Lozouet
<emphasis id="B90EEADB1938FFF3FEE1FDB2FE67F855" box="[375,422,576,600]" italics="true" pageId="14" pageNumber="259">et al.</emphasis>
2001: 45
</bibRefCitation>
</treatmentCitation>
, pl. 19, figs 3a-3b. —
<treatmentCitation id="0ADB10D81938FFF3FD4FFDB2FE84F87F" author="BEU A. G." page="68" pageId="14" pageNumber="259" year="2005">
<bibRefCitation id="EFEB4B381938FFF3FD4FFDB2FE84F87F" pageId="14" pageNumber="259" refId="ref11668" type="journal article">
Lan- dau
<emphasis id="B90EEADB1938FFF3FF39FDA9FF1FF87F" box="[175,222,602,626]" italics="true" pageId="14" pageNumber="259">et al.</emphasis>
2004: 68
</bibRefCitation>
</treatmentCitation>
. TYPE LOCALITY. —
<taxonomicName id="4C7A4D4A1938FFF3FED0FD78FE23F8AF" authorityName="Sanders &amp; Merle &amp; Puillandre" authorityYear="2019" baseAuthorityName="Grateloup" baseAuthorityYear="1833" box="[326,482,650,674]" class="Gastropoda" family="Ranellidae" genus="Ranella" kingdom="Animalia" order="Littorinimorpha" pageId="14" pageNumber="259" phylum="Mollusca" rank="species" species="tuberosa">
<emphasis id="B90EEADB1938FFF3FED0FD78FE23F8AF" box="[326,482,650,674]" italics="true" pageId="14" pageNumber="259">Ranella tuberosa</emphasis>
</taxonomicName>
and
<emphasis id="B90EEADB1938FFF3FD83FD78FD10F8AF" box="[533,721,650,674]" italics="true" pageId="14" pageNumber="259">Apollon pelouatensis</emphasis>
both from the Aquitanian of Saint-Paul-Lès-Dax,
<collectingCountry id="F36D76591938FFF3FDA1FD57FDBAF8B1" box="[567,635,677,700]" name="France" pageId="14" pageNumber="259">France</collectingCountry>
. TYPE MATERIAL. —
<taxonomicName id="4C7A4D4A1938FFF3FEDAFD26FE2BF8E1" authorityName="Sanders &amp; Merle &amp; Puillandre" authorityYear="2019" baseAuthorityName="Grateloup" baseAuthorityYear="1833" box="[332,490,724,748]" class="Gastropoda" family="Ranellidae" genus="Ranella" kingdom="Animalia" order="Littorinimorpha" pageId="14" pageNumber="259" phylum="Mollusca" rank="species" species="tuberosa">
<emphasis id="B90EEADB1938FFF3FEDAFD26FE2BF8E1" box="[332,490,724,748]" italics="true" pageId="14" pageNumber="259">Ranella tuberosa</emphasis>
</taxonomicName>
<typeStatus id="54C1886B1938FFF3FE64FD26FD87F8E1" box="[498,582,724,748]" pageId="14" pageNumber="259" type="syntype">syntypes</typeStatus>
in Grateloup coll., University of Bordeaux, Talence,
<collectingCountry id="F36D76591938FFF3FE52FD1DFDC4F90B" box="[452,517,751,774]" name="France" pageId="14" pageNumber="259">France</collectingCountry>
(not seen);
<emphasis id="B90EEADB1938FFF3FDE3FD1CFF74F92D" italics="true" pageId="14" pageNumber="259">Apollon peloua- tensis</emphasis>
: the specimen MNHN.F.
<accessionNumber id="9429AB2A1938FFF3FE22FCFBFE3DF92C" box="[436,508,777,801]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/J06123" pageId="14" pageNumber="259" type="EnaNcbi">J06123</accessionNumber>
is here designated as
<typeStatus id="54C1886B1938FFF3FD46FCFBFF70F931" pageId="14" pageNumber="259" type="lectotype">lectotype</typeStatus>
. Three
<typeStatus id="54C1886B1938FFF3FF6EFCD1FE43F936" box="[248,386,803,827]" pageId="14" pageNumber="259" type="paralectotype">paralectotypes</typeStatus>
(MNHN.F.
<accessionNumber id="9429AB2A1938FFF3FE6CFCD1FD82F936" box="[506,579,803,827]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/J06124" pageId="14" pageNumber="259" type="EnaNcbi">J06124</accessionNumber>
,
<accessionNumber id="9429AB2A1938FFF3FDD9FCD1FD56F936" box="[591,663,803,827]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/J06125" pageId="14" pageNumber="259" type="EnaNcbi">J06125</accessionNumber>
,
<accessionNumber id="9429AB2A1938FFF3FD32FCD1FD2DF936" box="[676,748,803,827]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/J06126" pageId="14" pageNumber="259" type="EnaNcbi">J06126</accessionNumber>
).. OTHER MATERIAL EXAMINED. —Two specimens from Hofstetter coll., from le Peloua, Burdigalian, and one from Saubrigues, Burdigalian;
<specimenCount id="9D7CFD401938FFF3FF12FC7AFEC9F9AD" box="[132,264,904,928]" pageId="14" pageNumber="259" type="generic">13 specimens</specimenCount>
from Brongniart coll., from le Peloua, Burdigalian;
<specimenCount id="9D7CFD401938FFF3FF12FC50FEC8F9B7" box="[132,265,930,954]" pageId="14" pageNumber="259" type="generic">15 specimens</specimenCount>
from Staadt coll., from le Peloua, Burdigalian; one specimen from Lhomme coll., from le Peloua, Burdigalian; two specimens (MNHN.F.
<accessionNumber id="9429AB2A1938FFF3FEF6FC25FE71F9E2" box="[352,432,983,1007]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/A70577" pageId="14" pageNumber="259" type="EnaNcbi">A70577</accessionNumber>
,
<accessionNumber id="9429AB2A1938FFF3FE2AFC25FDCDF9E2" box="[444,524,983,1007]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/A70578" pageId="14" pageNumber="259" type="EnaNcbi">A70578</accessionNumber>
) from Gaas “Lagouarde”, Rupelian; all in the collections de paléontologie, MNHN.
<subScript id="17FE348C1938FFF3FF12FBD3FDA0FE3A" attach="left" box="[132,609,1057,1081]" fontSize="7" pageId="14" pageNumber="259">GEOGRAPHIC AND STRATIGRAPHIC OCCURRENCE</subScript>
. — Rupelian to middle Miocene of the
<collectingRegion id="49BEF82B1938FFF3FECAFBCEFE78FE59" box="[348,441,1084,1108]" country="France" name="Aquitaine" pageId="14" pageNumber="259">Aquitaine</collectingRegion>
basin (
<collectingCountry id="F36D76591938FFF3FE6CFBCEFDFDFE59" box="[506,572,1084,1108]" name="France" pageId="14" pageNumber="259">France</collectingCountry>
) and Po Valley (
<collectingCountry id="F36D76591938FFF3FD58FBCEFD3CFE59" box="[718,765,1084,1108]" name="Italy" pageId="14" pageNumber="259">Italy</collectingCountry>
). RECOMMENDATIONS FOR CALIBRATION
</bibRefCitation>
</treatmentCitation>
</paragraph>
<paragraph id="8BC536C91938FFF3FF12FB24FAD3FBBF" blockId="14.[132,776,1174,2030]" lastBlockId="14.[813,1456,217,1359]" pageId="14" pageNumber="259">
In regard to the systematic considerations, we can provide some recommendations for calibration of the molecular tree of the
<taxonomicName id="4C7A4D4A1938FFF3FF12FAE7FF25FF22" ID-CoL="7NG76" authorityName="Thiele" authorityYear="1925" box="[132,228,1301,1327]" class="Gastropoda" family="Bursidae" kingdom="Animalia" order="Littorinimorpha" pageId="14" pageNumber="259" phylum="Mollusca" rank="family">Bursidae</taxonomicName>
. Numerical dating for this section is provided by the international geological time scale (
<bibRefCitation id="EFEB4B381938FFF3FD94FAC7FD24FF42" author="GRADSTEIN F. M. &amp; OGG J. G. &amp; SCHMITZ M. &amp; OGG G." box="[514,741,1333,1359]" pageId="14" pageNumber="259" refId="ref11131" refString="GRADSTEIN F. M., OGG J. G., SCHMITZ M. &amp; OGG G. 2012. - The Geologic Time Scale 2012. First Edition. Elsevier, New York, 1176 p." type="book" year="2012">
Gradstein
<emphasis id="B90EEADB1938FFF3FDE5FAC4FD67FF42" box="[627,678,1333,1359]" italics="true" pageId="14" pageNumber="259">et al.</emphasis>
2012
</bibRefCitation>
); a complete stratigraphic distribution chart for the
<taxonomicName id="4C7A4D4A1938FFF3FD04FAA7FD2EFF62" ID-CoL="7NG76" authorityName="Thiele" authorityYear="1925" box="[658,751,1365,1391]" class="Gastropoda" family="Bursidae" kingdom="Animalia" order="Littorinimorpha" pageId="14" pageNumber="259" phylum="Mollusca" rank="family">Bursidae</taxonomicName>
is shown in
<figureCitation id="13412A4C1938FFF3FF7BFA87FE87FF82" box="[237,326,1397,1423]" captionStart="FIG" captionStartId="15.[132,143,1941,1959]" captionTargetBox="[138,1450,215,1915]" captionTargetId="graphics@15.[139,1062,215,1914]" captionTargetPageId="15" captionText="FIG. 8. — Stratigraphic range of the extinct and extant Bursidae. Black arrows indicate total stratigraphic range; grey arrows indicate regional stratigraphic range.Abbreviations:P, Peru; WI, Western Indian Ocean; AB, Aquitaine Basin; CWA, Caribbean, Western Atlantic Ocean; PT, Paratethys, CWT: Central Western Tethys; EA, Eastern Atlantic; CWM, Central Western Mediterranean; WP, Western Pacific Ocean; J, Java, Indo-Pacific; S, Sumatra, Indo-Pacific; EP, Eastern Pacific Ocean. Grey squares regroup accepted genera and putative genera within Bursa." figureDoi="http://doi.org/10.5281/zenodo.3699948" httpUri="https://zenodo.org/record/3699948/files/figure.png" pageId="14" pageNumber="259">Figure 8</figureCitation>
. First occurrence and last occurrence data are provided in Appendix 1.
<taxonomicName id="4C7A4D4A1938FFF3FF0DFA47FAD3FBBF" authority="Considering" authorityName="Considering" class="Gastropoda" family="Bursidae" kingdom="Animalia" order="Littorinimorpha" pageId="14" pageNumber="259" phylum="Mollusca" rank="family">
Considering that it is the oldest undisputable taxon included in Bursidae and apparently is not closely related to other clades within the family, the genus
<emphasis id="B90EEADB1938FFF3FDCBFA07FD76FC02" box="[605,695,1525,1551]" italics="true" pageId="14" pageNumber="259">Olssonia</emphasis>
<taxonomicNameLabel id="A23D57A01938FFF3FD2BFA07FCC6FC02" box="[701,775,1525,1551]" pageId="14" pageNumber="259" rank="genus">n. gen.</taxonomicNameLabel>
seems to be the best candidate for the calibration of the whole Bursidae. The oldest representative of the genus is
<emphasis id="B90EEADB1938FFF3FF12F9A7FF23FC63" box="[132,226,1620,1647]" italics="true" pageId="14" pageNumber="259">O. yasila</emphasis>
<taxonomicNameLabel id="A23D57A01938FFF3FF7FF9A7FE8DFC63" box="[233,332,1620,1647]" pageId="14" pageNumber="259" rank="species">n. comb.</taxonomicNameLabel>
from the early Bartonian (
<bibRefCitation id="EFEB4B381938FFF3FDE5F9A6FCC1FC63" author="OLSSON A. A." box="[627,768,1620,1646]" pageId="14" pageNumber="259" pagination="1 - 272" refId="ref12299" refString="OLSSON A. A. 1932. - Contributions to the Tertiary paleontology of northern Peru: Part 5. The Peruvian Miocene. Bulletins of American Paleontology 19 (68): 1 - 272. https: // biodiversitylibrary. org / page / 32532230" type="journal article" year="1932">Olsson 1932</bibRefCitation>
,
<bibRefCitation id="EFEB4B381938FFF3FF12F987FF2EFC83" author="BEU A. G." box="[132,239,1652,1679]" pageId="14" pageNumber="259" pagination="69 - 96" refId="ref10328" refString="BEU A. G. 1988. - Taxonomy of gastropods of the families Ranellidae (= Cymatiidae) and Bursidae. Part 5. Early history of the families, with four new genera and recognition of the Family Personidae, in GRANT- MACKIE J. A., MASUDA K., MORI K. &amp; OGASAWARA K. (eds), ProfessorTamio Kotaka CommemorativeVolume on Molluscan Paleontology. Saito Ho-on Kai, Special Publication No. 2, Sendai: 69 - 96." type="book chapter" year="1988">Beu 1988</bibRefCitation>
) for which we propose to use an age of 41.2 Ma. As for all the following node calibrations, we propose to apply a truncated log normal distribution, as it places the highest probability on ages somewhat older than the fossil, diversification being necessarily older than the observed fossil record (
<bibRefCitation id="EFEB4B381938FFF3FE8FF8E6FE36FD23" author="HO S. Y. W. &amp; PHILLIPS M. J." box="[281,503,1812,1838]" pageId="14" pageNumber="259" pagination="367 - 380" refId="ref11538" refString="HO S. Y. W. &amp; PHILLIPS M. J. 2009. - Accounting for calibration uncertainty in phylogenetic estimation of evolutionary divergence times. Systematic Biology 58 (3): 367 - 380. https: // doi. org / 10.1093 / sysbio / syp 035" type="journal article" year="2009">Ho &amp; Phillips 2009</bibRefCitation>
). The genus
<emphasis id="B90EEADB1938FFF3FE9FF8C6FEF8FD43" box="[265,313,1844,1870]" italics="true" pageId="14" pageNumber="259">Aspa</emphasis>
does not resemble any other Bursidae closely and, considering its very long stratigraphic history, possibly commencing before the Neogene (
<bibRefCitation id="EFEB4B381938FFF3FE78F881FD71FD80" author="LANDAU B. &amp; BEU A. G. &amp; MARQUET R." box="[494,688,1907,1933]" pageId="14" pageNumber="259" pagination="35 - 102" refId="ref11668" refString="LANDAU B., BEU A. G. &amp; MARQUET R. 2004. - The Early Pliocene Gastropoda (Mollusca) of Estepona, Southern Spain. Part 5: Tonnoidea, Ficoidea. Palaeontos 5: 35 - 102." type="journal article" year="2004">
Landau
<emphasis id="B90EEADB1938FFF3FDD5F886FDB5FD80" box="[579,628,1907,1933]" italics="true" pageId="14" pageNumber="259">et al.</emphasis>
2004
</bibRefCitation>
), it may very well be a sister group to the rest of the Recent Bursidae. On the other hand, the fossil genus
<emphasis id="B90EEADB1938FFF3FE7BF841FD41FDC0" box="[493,640,1971,1997]" italics="true" pageId="14" pageNumber="259">Aquitanobursa</emphasis>
<taxonomicNameLabel id="A23D57A01938FFF3FD10F846FD0DFDC3" box="[646,716,1972,1998]" pageId="14" pageNumber="259" rank="genus">n. gen.</taxonomicNameLabel>
looks like some Recent species such as
<emphasis id="B90EEADB1938FFF3FE4AF826FE2DFDE0" box="[476,492,2004,2029]" italics="true" pageId="14" pageNumber="259">B</emphasis>
.
<emphasis id="B90EEADB1938FFF3FE6FF821FDA2FDE0" box="[505,611,2003,2029]" italics="true" pageId="14" pageNumber="259">quirihorai</emphasis>
and
<emphasis id="B90EEADB1938FFF3FD00F826FD67FDE0" box="[662,678,2004,2029]" italics="true" pageId="14" pageNumber="259">B</emphasis>
.
<emphasis id="B90EEADB1938FFF3FD25F826FCC9FDE0" box="[691,776,2004,2029]" italics="true" pageId="14" pageNumber="259">granosa.</emphasis>
If
<emphasis id="B90EEADB1938FFF3FCD0FF28FCB6FAF9" box="[838,887,218,244]" italics="true" pageId="14" pageNumber="259">Aspa</emphasis>
were to branch at the base of the Bursidae tree, the clade containing every other bursid but excluding
<emphasis id="B90EEADB1938FFF3FAABFF0BFAAFFB1E" box="[1341,1390,249,275]" italics="true" pageId="14" pageNumber="259">Aspa</emphasis>
could be calibrated by
<emphasis id="B90EEADB1938FFF3FC4FFEEBFBB3FB3E" box="[985,1138,281,307]" italics="true" pageId="14" pageNumber="259">Aquitanobursa</emphasis>
<taxonomicNameLabel id="A23D57A01938FFF3FBEEFEE8FB00FB39" box="[1144,1217,282,308]" pageId="14" pageNumber="259" rank="genus">n. gen.</taxonomicNameLabel>
The oldest representa- tive of the genus (two specimens MNHN.F.
<accessionNumber id="9429AB2A1938FFF3FB66FECBFA84FB59" box="[1264,1349,313,340]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/A70577" pageId="14" pageNumber="259" type="EnaNcbi">A70577</accessionNumber>
,
<accessionNumber id="9429AB2A1938FFF3FAC7FECBFA67FB59" box="[1361,1446,313,340]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/A70578" pageId="14" pageNumber="259" type="EnaNcbi">A70578</accessionNumber>
) is
<emphasis id="B90EEADB1938FFF3FCD2FEABFC76FB7E" box="[836,951,345,371]" italics="true" pageId="14" pageNumber="259">A. tuberosa</emphasis>
<taxonomicNameLabel id="A23D57A01938FFF3FC28FEABFBDEFB7E" box="[958,1055,345,371]" pageId="14" pageNumber="259" rank="species">n. comb.</taxonomicNameLabel>
from the Rupelian (Stampian) of the
<collectingRegion id="49BEF82B1938FFF3FCBBFE8BFC56FB9E" box="[813,919,377,403]" country="France" name="Aquitaine" pageId="14" pageNumber="259">Aquitaine</collectingRegion>
Basin. We propose an age of 27.82 Ma (Rupelian/ Chattian transition) for the node calibration.
</taxonomicName>
</paragraph>
<paragraph id="8BC536C91938FFF3FCD5FE4BFAEDF8BF" blockId="14.[813,1456,217,1359]" pageId="14" pageNumber="259">
<emphasis id="B90EEADB1938FFF3FCD5FE4BFC45FBDF" box="[835,900,441,466]" italics="true" pageId="14" pageNumber="259">
<taxonomicName id="4C7A4D4A1938FFF3FCD5FE4BFC41FBDF" ID-CoL="7NRCN" box="[835,896,441,466]" class="Gastropoda" family="Bursidae" genus="Bursa" kingdom="Animalia" order="Littorinimorpha" pageId="14" pageNumber="259" phylum="Mollusca" rank="genus">Bursa</taxonomicName>
,
</emphasis>
under the definition suggested in this paper, could be calibrated by the specimen resembling
<taxonomicName id="4C7A4D4A1938FFF3FB2FFE2BFB3AFBFF" ID-CoL="NY85" baseAuthorityName="Perry" baseAuthorityYear="1811" box="[1209,1275,473,498]" class="Gastropoda" family="Bursidae" genus="Bursa" kingdom="Animalia" order="Littorinimorpha" pageId="14" pageNumber="259" phylum="Mollusca" rank="species" species="rosa">
<emphasis id="B90EEADB1938FFF3FB2FFE2BFB3AFBFF" box="[1209,1275,473,498]" italics="true" pageId="14" pageNumber="259">B. rosa</emphasis>
</taxonomicName>
from middle-late Miocene of Citalahab,
<collectingRegion id="49BEF82B1938FFF3FBB0FE0BFB4FF81E" box="[1062,1166,505,531]" country="Indonesia" name="Jawa Barat" pageId="14" pageNumber="259">West Java</collectingRegion>
mentioned by Beu (2005: fig. 33). He attributed this specimen to
<emphasis id="B90EEADB1938FFF3FB4CFDEBFAE8F83F" box="[1242,1321,537,562]" italics="true" pageId="14" pageNumber="259">
<taxonomicName id="4C7A4D4A1938FFF3FB4CFDEBFAE7F83F" ID-CoL="NY85" baseAuthorityName="Perry" baseAuthorityYear="1811" box="[1242,1318,537,562]" class="Gastropoda" family="Bursidae" genus="Bursa" kingdom="Animalia" order="Littorinimorpha" pageId="14" pageNumber="259" phylum="Mollusca" rank="species" species="rosa">B. rosa</taxonomicName>
.
</emphasis>
Indeed, this specimen bears the long tubular posterior siphonal canal characteristic of the genus, but it is far less warty and could warrant erecting a new species, becoming thus the oldest spe- cies of all
<taxonomicName id="4C7A4D4A1938FFF3FC01FD6AFC13F8BC" ID-CoL="7NRCN" box="[919,978,664,689]" class="Gastropoda" family="Bursidae" genus="Bursa" kingdom="Animalia" order="Littorinimorpha" pageId="14" pageNumber="259" phylum="Mollusca" rank="genus" sensu="stricto">
<emphasis id="B90EEADB1938FFF3FC01FD6AFC13F8BC" box="[919,978,664,689]" italics="true" pageId="14" pageNumber="259">Bursa</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A23D57A01938FFF3FC4FFD6AFC37F8BF" box="[985,1014,664,690]" pageId="14" pageNumber="259" sensu="stricto">
<emphasis id="B90EEADB1938FFF3FC4FFD6AFC2FF8BC" box="[985,1006,664,689]" italics="true" pageId="14" pageNumber="259">s.s</emphasis>
.
</taxonomicNameLabel>
We propose an age of 9 Ma.
</paragraph>
<paragraph id="8BC536C91938FFF3FCD5FD45FBCDFE9C" blockId="14.[813,1456,217,1359]" pageId="14" pageNumber="259">
All the species of the former
<taxonomicName id="4C7A4D4A1938FFF3FB05FD4AFA86F8DC" ID-CoL="7QH9Z" baseAuthorityName="Roding" baseAuthorityYear="1798" box="[1171,1351,695,721]" class="Gastropoda" family="Bursidae" genus="Bursa" kingdom="Animalia" order="Littorinimorpha" pageId="14" pageNumber="259" phylum="Mollusca" rank="species" species="granularis">
<emphasis id="B90EEADB1938FFF3FB05FD4AFA86F8DC" box="[1171,1351,695,721]" italics="true" pageId="14" pageNumber="259">Bursa granularis</emphasis>
</taxonomicName>
complex (
<bibRefCitation id="EFEB4B381938FFF3FCA0FD25FC3EF8FF" author="SANDERS M. T. &amp; MERLE D. &amp; BOUCHET P. &amp; CASTELIN M. &amp; BEU A. G. &amp; SAMADI S. &amp; PUILLANDRE N." box="[822,1023,727,754]" pageId="14" pageNumber="259" pagination="384 - 398" refId="ref12735" refString="SANDERS M. T., MERLE D., BOUCHET P., CASTELIN M., BEU A. G., SAMADI S. &amp; PUILLANDRE N. 2017. - One for each ocean: revision of the Bursa granularis species complex (Gastropoda: Tonnoidea: Bursidae). Journal of Molluscan Studies 83 (4): 384 - 398. https: // doi. org / 10.1093 / mollus / eyx 029" type="journal article" year="2017">
Sanders
<emphasis id="B90EEADB1938FFF3FC19FD2AFC03F8FC" box="[911,962,727,753]" italics="true" pageId="14" pageNumber="259">et al.</emphasis>
2017
</bibRefCitation>
) could have their clade calibrated by the three specimens from Citalahab,
<collectingRegion id="49BEF82B1938FFF3FB1FFD0AFB2EF91F" box="[1161,1263,760,786]" country="Indonesia" name="Jawa Barat" pageId="14" pageNumber="259">West Java</collectingRegion>
attributed by Beu (2005) to
<taxonomicName id="4C7A4D4A1938FFF3FC0DFCEAFBE0F93C" ID-CoL="7QH9Z" baseAuthorityName="Roding" baseAuthorityYear="1798" box="[923,1057,791,817]" class="Gastropoda" family="Bursidae" genus="Bursa" kingdom="Animalia" order="Littorinimorpha" pageId="14" pageNumber="259" phylum="Mollusca" rank="species" species="granularis">
<emphasis id="B90EEADB1938FFF3FC0DFCEAFBE0F93C" box="[923,1057,791,817]" italics="true" pageId="14" pageNumber="259">B. granularis</emphasis>
</taxonomicName>
, with an age of 9 Ma as for
<emphasis id="B90EEADB1938FFF3FAC6FCEAFA71F93C" box="[1360,1456,792,817]" italics="true" pageId="14" pageNumber="259">
<taxonomicName id="4C7A4D4A1938FFF3FAC6FCEAFA4AF93C" ID-CoL="7NRCN" box="[1360,1419,792,817]" class="Gastropoda" family="Bursidae" genus="Bursa" kingdom="Animalia" order="Littorinimorpha" pageId="14" pageNumber="259" phylum="Mollusca" rank="genus" sensu="stricto">Bursa</taxonomicName>
<taxonomicNameLabel id="A23D57A01938FFF3FA04FCEAFA71F93C" box="[1426,1456,792,817]" pageId="14" pageNumber="259" sensu="stricto">s.s.</taxonomicNameLabel>
</emphasis>
This calibration should not be blindly trusted, because we have not seen the specimens and they were not figured by Beu (2005). The relationship of
<emphasis id="B90EEADB1938FFF3FBCFFC8AFB08F99C" box="[1113,1225,887,913]" italics="true" pageId="14" pageNumber="259">B. landaui</emphasis>
to this clade remains too uncertain to be used as a calibration. The genus
<taxonomicName id="4C7A4D4A1938FFF3FC22FC45FBE7F9DD" ID-CoL="7P8HH" authorityName="Dall" authorityYear="1904" box="[948,1062,951,976]" class="Gastropoda" family="Bursidae" genus="Marsupina" kingdom="Animalia" order="Littorinimorpha" pageId="14" pageNumber="259" phylum="Mollusca" rank="genus">
<emphasis id="B90EEADB1938FFF3FC22FC45FBE7F9DD" box="[948,1062,951,976]" italics="true" pageId="14" pageNumber="259">Marsupina</emphasis>
</taxonomicName>
could be calibrated by the Shoal river formation specimen (USNM 647109), middle Miocene, fig- ured by
<bibRefCitation id="EFEB4B381938FFF3FC17FC05FC30FE1C" author="BEU A. G." box="[897,1009,1015,1041]" pageId="14" pageNumber="259" pagination="1 - 550" refId="ref10501" refString="BEU A. G. 2010. - Neogene tonnoidean gastropods of tropical and South America; a contribution to the Dominican Republic and Panama Paleontology projects and uplift of the Central American Isthmus. Bulletins of American Paleontology 377 - 378: 1 - 550." type="journal article" year="2010">Beu (2010</bibRefCitation>
: pl. 9, fig. 6). This juvenile specimen strik- ingly resembling
<emphasis id="B90EEADB1938FFF3FC75FBE5FBFCFE3D" box="[995,1085,1046,1072]" italics="true" pageId="14" pageNumber="259">
<taxonomicName id="4C7A4D4A1938FFF3FC75FBE5FBF8FE3D" ID-CoL="3Y9PM" baseAuthorityName="Bruguière" baseAuthorityYear="1792" box="[995,1081,1046,1072]" class="Gastropoda" family="Bursidae" genus="Marsupina" kingdom="Animalia" order="Littorinimorpha" pageId="14" pageNumber="259" phylum="Mollusca" rank="species" species="bufo">M. bufo</taxonomicName>
,
</emphasis>
but has a more coarsely sculptured shell than later specimens of
<taxonomicName id="4C7A4D4A1938FFF3FBF1FBC5FB18FE5D" ID-CoL="7P8HH" authorityName="Dall" authorityYear="1904" box="[1127,1241,1079,1104]" class="Gastropoda" family="Bursidae" genus="Marsupina" kingdom="Animalia" order="Littorinimorpha" pageId="14" pageNumber="259" phylum="Mollusca" rank="genus">
<emphasis id="B90EEADB1938FFF3FBF1FBC5FB18FE5D" box="[1127,1241,1079,1104]" italics="true" pageId="14" pageNumber="259">Marsupina</emphasis>
</taxonomicName>
, too much so to be conspecific. We propose an age of 14 Ma for the calibration of crown
<taxonomicName id="4C7A4D4A1938FFF3FC05FB85FBC4FE9D" ID-CoL="7P8HH" authorityName="Dall" authorityYear="1904" box="[915,1029,1143,1168]" class="Gastropoda" family="Bursidae" genus="Marsupina" kingdom="Animalia" order="Littorinimorpha" pageId="14" pageNumber="259" phylum="Mollusca" rank="genus">
<emphasis id="B90EEADB1938FFF3FC05FB85FBC4FE9D" box="[915,1029,1143,1168]" italics="true" pageId="14" pageNumber="259">Marsupina</emphasis>
</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection id="C36065421938FFEDFCD5FB64FD28FCE3" lastPageId="16" lastPageNumber="261" pageId="14" pageNumber="259" type="description">
<paragraph id="8BC536C91938FFF3FCD5FB64FAE0FDE0" blockId="14.[813,1456,217,1359]" lastBlockId="14.[813,1456,1429,2029]" pageId="14" pageNumber="259">
Strong
<emphasis id="B90EEADB1938FFF3FC04FB65FC07FEBD" box="[914,966,1174,1200]" italics="true" pageId="14" pageNumber="259">et al.</emphasis>
(2019) proposed to use the large specimen of
<emphasis id="B90EEADB1938FFF3FCBBFB44FCB2FEDD" box="[813,883,1206,1232]" italics="true" pageId="14" pageNumber="259">Tutufa</emphasis>
sp. Looking a lot like
<emphasis id="B90EEADB1938FFF3FBC8FB44FB6CFEDD" box="[1118,1197,1206,1232]" italics="true" pageId="14" pageNumber="259">T. bubo</emphasis>
figured by Tomida
<emphasis id="B90EEADB1938FFF3FAEBFB44FA71FEDD" box="[1405,1456,1206,1232]" italics="true" pageId="14" pageNumber="259">et al.</emphasis>
(2013) as a calibration point for the genus
<emphasis id="B90EEADB1938FFF3FB65FB24FAF8FEFD" box="[1267,1337,1238,1264]" italics="true" pageId="14" pageNumber="259">Tutufa</emphasis>
, calibrated at 13.5 Ma. We follow their recommendation. No other node could be calibrated easily based on what we know currently of the systematics of
<taxonomicName id="4C7A4D4A1938FFF3FB2FFAC7FADBFF42" authorityName="Thiele" authorityYear="1925" box="[1209,1306,1333,1359]" class="Gastropoda" family="Bursidae" kingdom="Animalia" order="Littorinimorpha" pageId="14" pageNumber="259" phylum="Mollusca" rank="family">Bursidae</taxonomicName>
. TIMING OF DIVERSIFICATION OF BURSIDAE BASED ON THE FOSSIL RECORD Providing a scenario explaining the diversification of a highly dispersive family like the
<taxonomicName id="4C7A4D4A1938FFF3FBD7F9E6FB5FFC23" box="[1089,1182,1556,1582]" class="Gastropoda" family="Bursidae" kingdom="Animalia" order="Littorinimorpha" pageId="14" pageNumber="259" phylum="Mollusca" rank="family">Bursidae</taxonomicName>
is not an easy task, especially without a proper phylogenetic context. What we can say, however, is that the diversification of the
<taxonomicName id="4C7A4D4A1938FFF3FA8CF9A6FAB6FC63" box="[1306,1399,1620,1646]" class="Gastropoda" family="Bursidae" kingdom="Animalia" order="Littorinimorpha" pageId="14" pageNumber="259" phylum="Mollusca" rank="family">Bursidae</taxonomicName>
hap- pened in three phases (
<figureCitation id="13412A4C1938FFF3FBB2F986FBA3FC83" box="[1060,1122,1652,1678]" captionStart="FIG" captionStartId="15.[132,143,1941,1959]" captionTargetBox="[138,1450,215,1915]" captionTargetId="graphics@15.[139,1062,215,1914]" captionTargetPageId="15" captionText="FIG. 8. — Stratigraphic range of the extinct and extant Bursidae. Black arrows indicate total stratigraphic range; grey arrows indicate regional stratigraphic range.Abbreviations:P, Peru; WI, Western Indian Ocean; AB, Aquitaine Basin; CWA, Caribbean, Western Atlantic Ocean; PT, Paratethys, CWT: Central Western Tethys; EA, Eastern Atlantic; CWM, Central Western Mediterranean; WP, Western Pacific Ocean; J, Java, Indo-Pacific; S, Sumatra, Indo-Pacific; EP, Eastern Pacific Ocean. Grey squares regroup accepted genera and putative genera within Bursa." figureDoi="http://doi.org/10.5281/zenodo.3699948" httpUri="https://zenodo.org/record/3699948/files/figure.png" pageId="14" pageNumber="259">Fig. 8</figureCitation>
).
<bibRefCitation id="EFEB4B381938FFF3FCD5F966FBE9FCA2" author="STRONG E. E. &amp; PUILLANDRE N. &amp; BEU A. G. &amp; CASTELIN M. &amp; BOUCHET P." box="[835,1064,1684,1711]" pageId="14" pageNumber="259" pagination="18 - 34" refId="ref13233" refString="STRONG E. E., PUILLANDRE N., BEU A. G., CASTELIN M. &amp; BOUCHET P. 2018. - Frogs and tuns and tritons - A molecular phylogeny and revised family classification of the predatory gastropod superfamily Tonnoidea (Caenogastropoda). Molecular Phylogenetics and Evolution 130: 18 - 34. https: // doi. org / 10.1016 / j. ympev. 2018.09.016" type="journal article" year="2018">
Strong
<emphasis id="B90EEADB1938FFF3FC0EF967FC10FCA3" box="[920,977,1684,1710]" italics="true" pageId="14" pageNumber="259">et al.</emphasis>
(2018)
</bibRefCitation>
estimated (with very low support regarding its relationship with its sister family) the origin of the family at around 113 Ma but, as stated above, we have no trace of morphologically recognizable bursids prior to the middle Eocene (40 Ma) Peruvian fauna. Following this record there is no other trace of
<taxonomicName id="4C7A4D4A1938FFF3FBE9F8C1FB1DFD40" box="[1151,1244,1843,1869]" class="Gastropoda" family="Bursidae" kingdom="Animalia" order="Littorinimorpha" pageId="14" pageNumber="259" phylum="Mollusca" rank="family">Bursidae</taxonomicName>
in the Paleogene. A possible explanation of this lack of fossils is a general diminution of fossil-rich (unlithified) marine outcrops in the late Paleogene (
<bibRefCitation id="EFEB4B381938FFF3FC3EF861FBF3FDA0" author="HENDY A. J. W." box="[936,1074,1939,1965]" pageId="14" pageNumber="259" pagination="19 - 77" refId="ref11464" refString="HENDY A. J. W. 2011. - Taphonomic Overprints on Phanerozoic Trends in Biodiversity: Lithification and Other Secular Megabiases, in ALLISON P. A. &amp; BOTTJER D. J. (eds), Taphonomy. Aims &amp; Scope Topics in Geobiology 32: 19 - 77. https: // doi. org / 10.1007 / 978 - 90 - 481 - 8643 - 3 _ 2" type="journal article" year="2011">Hendy 2011</bibRefCitation>
). This tendency dramatically shifts in the Neogene, as we found an already well diversified fauna throughout the lower Miocene Tethyan realm.
</paragraph>
<caption id="DF0566411939FFF2FF12F867FCACFDE6" ID-DOI="http://doi.org/10.5281/zenodo.3699948" ID-Zenodo-Dep="3699948" httpUri="https://zenodo.org/record/3699948/files/figure.png" pageId="15" pageNumber="260" startId="15.[132,143,1941,1959]" targetBox="[138,1450,215,1915]" targetPageId="15">
<paragraph id="8BC536C91939FFF2FF12F867FCACFDE6" blockId="15.[132,1456,1941,2028]" pageId="15" pageNumber="260">
FIG. 8. — Stratigraphic range of the extinct and extant
<taxonomicName id="4C7A4D4A1939FFF2FDC3F867FD63FDAA" box="[597,674,1941,1959]" class="Gastropoda" family="Bursidae" kingdom="Animalia" order="Littorinimorpha" pageId="15" pageNumber="260" phylum="Mollusca" rank="family">Bursidae</taxonomicName>
.
<emphasis id="B90EEADB1939FFF2FD3DF864FCE0FDA5" bold="true" box="[683,801,1942,1960]" pageId="15" pageNumber="260">Black arrows</emphasis>
indicate total stratigraphic range;
<emphasis id="B90EEADB1939FFF2FBD3F864FB71FDA5" bold="true" box="[1093,1200,1942,1960]" pageId="15" pageNumber="260">grey arrows</emphasis>
indicate regional stratigraphic range.Abbreviations:
<emphasis id="B90EEADB1939FFF2FEAFF85EFE84FDB3" bold="true" box="[313,325,1964,1982]" pageId="15" pageNumber="260">P</emphasis>
, Peru;
<emphasis id="B90EEADB1939FFF2FE16F85EFE56FDB3" bold="true" box="[384,407,1964,1982]" pageId="15" pageNumber="260">WI</emphasis>
, Western Indian Ocean;
<emphasis id="B90EEADB1939FFF2FDF4F85EFDBDFDB3" bold="true" box="[610,636,1964,1982]" pageId="15" pageNumber="260">AB</emphasis>
, Aquitaine Basin;
<emphasis id="B90EEADB1939FFF2FC87F85EFCFDFDB3" bold="true" box="[785,828,1964,1982]" pageId="15" pageNumber="260">CWA</emphasis>
, Caribbean, Western Atlantic Ocean;
<emphasis id="B90EEADB1939FFF2FBE2F85EFB4DFDB3" bold="true" box="[1140,1164,1964,1982]" pageId="15" pageNumber="260">PT</emphasis>
, Paratethys, CWT: Central Western Tethys;
<emphasis id="B90EEADB1939FFF2FF53F831FF1FFDD8" bold="true" box="[197,222,1987,2005]" pageId="15" pageNumber="260">EA</emphasis>
, Eastern Atlantic;
<emphasis id="B90EEADB1939FFF2FEEFF831FE68FDD8" bold="true" box="[377,425,1987,2005]" pageId="15" pageNumber="260">CWM</emphasis>
, Central Western Mediterranean;
<emphasis id="B90EEADB1939FFF2FD50F831FD25FDD8" bold="true" box="[710,740,1987,2005]" pageId="15" pageNumber="260">WP</emphasis>
, Western Pacific Ocean;
<emphasis id="B90EEADB1939FFF2FC2DF831FC04FDD8" bold="true" box="[955,965,1987,2005]" pageId="15" pageNumber="260">J</emphasis>
, Java, Indo-Pacific;
<emphasis id="B90EEADB1939FFF2FBE4F831FBBFFDD8" bold="true" box="[1138,1150,1987,2005]" pageId="15" pageNumber="260">S</emphasis>
, Sumatra, Indo-Pacific;
<emphasis id="B90EEADB1939FFF2FADAF831FAA4FDD8" bold="true" box="[1356,1381,1987,2005]" pageId="15" pageNumber="260">EP</emphasis>
, Eastern Pacific Ocean.
<emphasis id="B90EEADB1939FFF2FE94F82BFEB9FDE1" bold="true" box="[258,376,2009,2028]" pageId="15" pageNumber="260">Grey squares</emphasis>
regroup accepted genera and putative genera within
<taxonomicName id="4C7A4D4A1939FFF2FCA0F82BFCA8FDE6" box="[822,873,2009,2027]" class="Gastropoda" family="Bursidae" genus="Bursa" kingdom="Animalia" order="Littorinimorpha" pageId="15" pageNumber="260" phylum="Mollusca" rank="genus">Bursa</taxonomicName>
.
</paragraph>
</caption>
<paragraph id="8BC536C91926FFEDFF0DFF28FCC6F87F" blockId="16.[132,775,217,1774]" pageId="16" pageNumber="261">
In the west (Aquitaine Basin) most species of the genus
<taxonomicName id="4C7A4D4A1926FFEDFD5AFF28FF26FB1E" authority="Sanders &amp; Merle &amp; Puillandre, 2019" authorityName="Sanders &amp; Merle &amp; Puillandre" authorityYear="2019" class="Gastropoda" family="Bursidae" genus="Aquitanobursa" higherTaxonomySource="GBIF" kingdom="Animalia" order="Littorinimorpha" pageId="16" pageNumber="261" phylum="Mollusca" rank="genus" status="gen. nov.">
<emphasis id="B90EEADB1926FFEDFD5AFF28FF26FB1E" italics="true" pageId="16" pageNumber="261">Aquitanobursa</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A23D57A01926FFEDFF7AFF08FEF4FB19" box="[236,309,250,276]" pageId="16" pageNumber="261" rank="genus">n. gen.</taxonomicNameLabel>
,
<emphasis id="B90EEADB1926FFEDFEABFF0BFE32FB1E" box="[317,499,249,275]" italics="true" pageId="16" pageNumber="261">
<taxonomicName id="4C7A4D4A1926FFEDFEABFF0BFEACFB1E" authorityName="H. Adams &amp; A. Adams" authorityYear="1853" box="[317,365,249,275]" class="Gastropoda" family="Bursidae" genus="Aspa" kingdom="Animalia" order="Littorinimorpha" pageId="16" pageNumber="261" phylum="Mollusca" rank="genus">Aspa</taxonomicName>
subgranulata
</emphasis>
and
<taxonomicName id="4C7A4D4A1926FFEDFDB2FF08FD00FB1E" baseAuthorityName="Perry" baseAuthorityYear="1811" box="[548,705,250,275]" class="Gastropoda" family="Bursidae" genus="Bursa" kingdom="Animalia" order="Littorinimorpha" pageId="16" pageNumber="261" phylum="Mollusca" rank="species" species="corrugata">
<emphasis id="B90EEADB1926FFEDFDB2FF08FD00FB1E" box="[548,705,250,275]" italics="true" pageId="16" pageNumber="261">Bursa corrugata</emphasis>
</taxonomicName>
occur. In the “Mediterranean region” (
<emphasis id="B90EEADB1926FFEDFE42FEE8FDC6FB3E" box="[468,519,282,307]" italics="true" pageId="16" pageNumber="261">sensu</emphasis>
<bibRefCitation id="EFEB4B381926FFEDFD98FEEBFD3FFB3E" author="HARZHAUSER M. &amp; PILLER W. E. &amp; STEININGER F. F." box="[526,766,281,307]" pageId="16" pageNumber="261" pagination="103 - 133" refId="ref11275" refString="HARZHAUSER M., PILLER W. E. &amp; STEININGER F. F. 2002. - Circum- Mediterranean Oligo-Miocene biogeographic evolution - the gastropods' point of view. Palaeogeography, Palaeoclimatology, Palaeoecology 183 (1 - 2): 103 - 133. https: // doi. org / 10.1016 / S 0031 - 0182 (01) 00464 - 3" type="journal article" year="2002">
Harzhauser
<emphasis id="B90EEADB1926FFEDFD18FEE8FD01FB3E" box="[654,704,281,307]" italics="true" pageId="16" pageNumber="261">et al.</emphasis>
2002
</bibRefCitation>
)
<taxonomicName id="4C7A4D4A1926FFEDFF12FEC8FEE6FB5E" baseAuthorityName="Perry" baseAuthorityYear="1811" box="[132,295,314,339]" class="Gastropoda" family="Bursidae" genus="Bursa" kingdom="Animalia" order="Littorinimorpha" pageId="16" pageNumber="261" phylum="Mollusca" rank="species" species="corrugata">
<emphasis id="B90EEADB1926FFEDFF12FEC8FEE6FB5E" box="[132,295,314,339]" italics="true" pageId="16" pageNumber="261">Bursa corrugata</emphasis>
</taxonomicName>
and
<taxonomicName id="4C7A4D4A1926FFEDFEC9FEC8FDCDFB5E" baseAuthorityName="Reeve" baseAuthorityYear="1844" box="[351,524,313,339]" class="Gastropoda" family="Bursidae" genus="Bursa" kingdom="Animalia" order="Littorinimorpha" pageId="16" pageNumber="261" phylum="Mollusca" rank="species" species="ranelloides">
<emphasis id="B90EEADB1926FFEDFEC9FEC8FDCDFB5E" box="[351,524,313,339]" italics="true" pageId="16" pageNumber="261">Bursa ranelloides</emphasis>
</taxonomicName>
occur. The presence of
<taxonomicName id="4C7A4D4A1926FFEDFF12FEABFEC0FB7F" baseAuthorityName="Perry" baseAuthorityYear="1811" box="[132,257,345,370]" class="Gastropoda" family="Bursidae" genus="Bursa" kingdom="Animalia" order="Littorinimorpha" pageId="16" pageNumber="261" phylum="Mollusca" rank="species" species="corrugata">
<emphasis id="B90EEADB1926FFEDFF12FEABFEC0FB7F" box="[132,257,345,370]" italics="true" pageId="16" pageNumber="261">B. corrugata</emphasis>
</taxonomicName>
in both regions is a clear indication of a connection between the westernmost Tethys and the Atlantic. In the Eastern Proto-Indo-West-Pacific region,
<taxonomicName id="4C7A4D4A1926FFEDFD80FE6AFDACFBBF" box="[534,621,408,434]" class="Gastropoda" family="Bursidae" kingdom="Animalia" order="Littorinimorpha" pageId="16" pageNumber="261" phylum="Mollusca" rank="family">Bursidae</taxonomicName>
are represented by
<taxonomicName id="4C7A4D4A1926FFEDFF35FE4AFE47FBDF" authorityName="Sanders &amp; Merle &amp; Puillandre" authorityYear="2019" baseAuthorityName="d'Archiac &amp; Haimes" baseAuthorityYear="1853" box="[163,390,440,466]" class="Gastropoda" family="Bursidae" genus="Aquitanobursa" higherTaxonomySource="GBIF" kingdom="Animalia" order="Littorinimorpha" pageId="16" pageNumber="261" phylum="Mollusca" rank="species" species="morrisi" status="comb. nov.">
<emphasis id="B90EEADB1926FFEDFF35FE4AFE47FBDF" box="[163,390,440,466]" italics="true" pageId="16" pageNumber="261">Aquitanobursa morrisi</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A23D57A01926FFEDFE1AFE4BFE28FBDF" box="[396,489,440,467]" pageId="16" pageNumber="261" rank="species">n. comb.</taxonomicNameLabel>
in the north (
<collectingCountry id="F36D76591926FFEDFDEEFE4AFD12FBDE" box="[632,723,440,467]" name="Pakistan" pageId="16" pageNumber="261">Pakistan</collectingCountry>
) and by
<emphasis id="B90EEADB1926FFEDFF30FE2BFEF6FBFF" box="[166,311,472,498]" italics="true" pageId="16" pageNumber="261">
<taxonomicName id="4C7A4D4A1926FFEDFF30FE2BFF21FBFF" box="[166,224,473,498]" class="Gastropoda" family="Bursidae" genus="Bursa" kingdom="Animalia" order="Littorinimorpha" pageId="16" pageNumber="261" phylum="Mollusca" rank="genus">Bursa</taxonomicName>
landaui
</emphasis>
in the southwest (
<collectingCountry id="F36D76591926FFEDFE69FE2AFDA5FBFE" box="[511,612,472,499]" name="Tanzania" pageId="16" pageNumber="261">Tanzania</collectingCountry>
). The arrival of
<taxonomicName id="4C7A4D4A1926FFEDFF12FE0BFF28F81F" authorityName="Sanders &amp; Merle &amp; Puillandre" authorityYear="2019" baseAuthorityName="d'Archiac &amp; Haimes" baseAuthorityYear="1853" box="[132,233,505,531]" class="Gastropoda" family="Bursidae" genus="Aquitanobursa" higherTaxonomySource="GBIF" kingdom="Animalia" order="Littorinimorpha" pageId="16" pageNumber="261" phylum="Mollusca" rank="species" species="morrisi" status="comb. nov.">
<emphasis id="B90EEADB1926FFEDFF12FE0BFF28F81F" box="[132,233,505,531]" italics="true" pageId="16" pageNumber="261">A. morrisi</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A23D57A01926FFEDFF79FE0BFE8CF81F" box="[239,333,504,531]" pageId="16" pageNumber="261" rank="species">n. comb.</taxonomicNameLabel>
in the Proto-Indo-West-Pacific region was most certainly through the Tethys, as most of the species of the genus are found in the Aquitaine Basin. On the other hand, the arrival
<emphasis id="B90EEADB1926FFEDFF65FDABFE9FF87F" box="[243,350,600,626]" italics="true" pageId="16" pageNumber="261">B. landaui</emphasis>
in the region is more difficult to explain.
</paragraph>
<paragraph id="8BC536C91926FFEDFF0DFD8AFD3CF99C" blockId="16.[132,775,217,1774]" pageId="16" pageNumber="261">
The next phase of diversification happened during the middle Miocene with the closure of the Tethys Ocean in the east. Following this closure, we observed a relocation of the main coral biodiversity hotspot from the Mediterranean to the present-day coral triangle (
<bibRefCitation id="EFEB4B381926FFEDFE5EFD05FD58F91C" author="LEPRIEUR F. &amp; DESCOMBES P. &amp; GABORIAU T. &amp; COWMAN P. F. &amp; PARRAVICINI V. &amp; KULBICKI M. &amp; MELIAN C. J. &amp; SANTANA C. N. DE &amp; HEINE C. &amp; MOUILLOT D. &amp; BELLWOOD D. R. &amp; PELLISSIER L." box="[456,665,759,785]" pageId="16" pageNumber="261" pagination="11461" refId="ref11799" refString="LEPRIEUR F., DESCOMBES P., GABORIAU T., COWMAN P. F., PARRAVICINI V., KULBICKI M., MELIAN C. J., SANTANA C. N. DE, HEINE C., MOUILLOT D., BELLWOOD D. R. &amp; PELLISSIER L. 2016. - Plate tectonics drive tropical reef biodiversity dynamics. Nature Communications 7: 11461. https: // doi. org / 10.1038 / ncomms 11461" type="journal article" year="2016">
Leprieur
<emphasis id="B90EEADB1926FFEDFDBCFD0AFD9DF91C" box="[554,604,759,785]" italics="true" pageId="16" pageNumber="261">et al.</emphasis>
2016
</bibRefCitation>
). This hot spot shift could explain the radiation of the coral-dependent bursids (
<emphasis id="B90EEADB1926FFEDFF75FCC5FEE8F95C" box="[227,297,823,849]" italics="true" pageId="16" pageNumber="261">Tutufa</emphasis>
,
<taxonomicName id="4C7A4D4A1926FFEDFEAFFCC5FE63F95C" box="[313,418,823,849]" class="Gastropoda" family="Bursidae" genus="Bufonaria" kingdom="Animalia" order="Littorinimorpha" pageId="16" pageNumber="261" phylum="Mollusca" rank="genus">
<emphasis id="B90EEADB1926FFEDFEAFFCC5FE63F95C" box="[313,418,823,849]" italics="true" pageId="16" pageNumber="261">Bufonaria</emphasis>
</taxonomicName>
,
<emphasis id="B90EEADB1926FFEDFE24FCCAFDD2F95C" box="[434,531,824,849]" italics="true" pageId="16" pageNumber="261">
<taxonomicName id="4C7A4D4A1926FFEDFE24FCCAFE2CF95C" box="[434,493,824,849]" class="Gastropoda" family="Bursidae" genus="Bursa" kingdom="Animalia" order="Littorinimorpha" pageId="16" pageNumber="261" phylum="Mollusca" rank="genus" sensu="stricto">Bursa</taxonomicName>
<taxonomicNameLabel id="A23D57A01926FFEDFE60FCCAFDD2F95C" box="[502,531,824,849]" pageId="16" pageNumber="261" sensu="stricto">s.s.</taxonomicNameLabel>
</emphasis>
). An eastward colonization of most of those species is probable, although some genera (e.g.
<emphasis id="B90EEADB1926FFEDFE91FC85FE4DF99C" box="[263,396,887,913]" italics="true" pageId="16" pageNumber="261">Lampadopsis</emphasis>
) possibly arrived from the Pacific.
</paragraph>
<paragraph id="8BC536C91926FFEDFF0DFC65FD01FE5D" blockId="16.[132,775,217,1774]" pageId="16" pageNumber="261">
In the west, also during the middle Miocene,
<taxonomicName id="4C7A4D4A1926FFEDFDFFFC65FCC6F9BD" baseAuthorityName="Gmelin" baseAuthorityYear="1791" box="[617,775,919,945]" class="Gastropoda" family="Bursidae" genus="Aspa" kingdom="Animalia" order="Littorinimorpha" pageId="16" pageNumber="261" phylum="Mollusca" rank="species" species="marginata">
<emphasis id="B90EEADB1926FFEDFDFFFC65FCC6F9BD" box="[617,775,919,945]" italics="true" pageId="16" pageNumber="261">Aspa marginata</emphasis>
</taxonomicName>
entered the Eastern Atlantic and
<emphasis id="B90EEADB1926FFEDFE58FC45FD51F9DD" box="[462,656,951,977]" italics="true" pageId="16" pageNumber="261">Lampadopsis rugosa</emphasis>
entered the Pacific Ocean from the Caribbean. The genus
<taxonomicName id="4C7A4D4A1926FFEDFDF8FC24FCC6F9FD" authority="Sanders &amp; Merle &amp; Puillandre, 2019" authorityName="Sanders &amp; Merle &amp; Puillandre" authorityYear="2019" box="[622,775,982,1008]" class="Gastropoda" family="Bursidae" genus="Aquitanobursa" higherTaxonomySource="GBIF" kingdom="Animalia" order="Littorinimorpha" pageId="16" pageNumber="261" phylum="Mollusca" rank="genus" status="gen. nov.">
<emphasis id="B90EEADB1926FFEDFDF8FC24FCC6F9FD" box="[622,775,982,1008]" italics="true" pageId="16" pageNumber="261">Aquitanobursa</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A23D57A01926FFEDFF12FC05FF0DFE1C" box="[132,204,1015,1041]" pageId="16" pageNumber="261" rank="genus">n. gen.</taxonomicNameLabel>
entered the Western Atlantic (
<taxonomicName id="4C7A4D4A1926FFEDFD9DFC05FD5AFE1D" authorityName="Sanders &amp; Merle &amp; Puillandre" authorityYear="2019" box="[523,667,1014,1041]" class="Gastropoda" family="Bursidae" genus="Bursa" kingdom="Animalia" order="Littorinimorpha" pageId="16" pageNumber="261" phylum="Mollusca" rank="species" species="amphitrites" status="comb. nov.">
<emphasis id="B90EEADB1926FFEDFD9DFC05FD5AFE1D" box="[523,667,1014,1041]" italics="true" pageId="16" pageNumber="261">A. amphitrites</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A23D57A01926FFEDFD36FC05FCC1FE1C" box="[672,768,1014,1041]" pageId="16" pageNumber="261" rank="species">n. comb.</taxonomicNameLabel>
) and the genus
<taxonomicName id="4C7A4D4A1926FFEDFE82FBE5FE40FE3D" authorityName="Dall" authorityYear="1904" box="[276,385,1047,1072]" class="Gastropoda" family="Bursidae" genus="Marsupina" kingdom="Animalia" order="Littorinimorpha" pageId="16" pageNumber="261" phylum="Mollusca" rank="genus">
<emphasis id="B90EEADB1926FFEDFE82FBE5FE40FE3D" box="[276,385,1047,1072]" italics="true" pageId="16" pageNumber="261">Marsupina</emphasis>
</taxonomicName>
appeared.In the Proto-Mediterranean Atlantic region the species
<taxonomicName id="4C7A4D4A1926FFEDFE33FBC5FD93FE5D" authorityName="" baseAuthorityName="Linnaeus" baseAuthorityYear="1758" box="[421,594,1078,1104]" class="Gastropoda" family="Bursidae" genus="Bursa" kingdom="Animalia" order="Littorinimorpha" pageId="16" pageNumber="261" phylum="Mollusca" rank="species" species="scrobilator">
<emphasis id="B90EEADB1926FFEDFE33FBC5FD93FE5D" box="[421,594,1078,1104]" italics="true" pageId="16" pageNumber="261">Bursa scrobilator</emphasis>
</taxonomicName>
appeared.
</paragraph>
<paragraph id="8BC536C91926FFEDFF0DFBA4FEF9FEBD" blockId="16.[132,775,217,1774]" pageId="16" pageNumber="261">
The high species-level diversity of tonnoideans in general during the middle Miocene was already pointed out by
<bibRefCitation id="EFEB4B381926FFEDFD42FB85FEF2FEBD" author="LANDAU B. &amp; HARZHAUSER M. &amp; BEU A. G." pageId="16" pageNumber="261" pagination="61 - 109" refId="ref11711" refString="LANDAU B., HARZHAUSER M. &amp; BEU A. G. 2009. - A Revision of the Tonnoidea (Caenogastropoda, Gastropoda) from the Miocene Paratethys and their palaeobiogeographic implications. Jahrbuch Der Geologischen Bundesanstalt 149: 61 - 109." type="journal article" year="2009">
Landau
<emphasis id="B90EEADB1926FFEDFF25FB65FF26FEBD" box="[179,231,1174,1200]" italics="true" pageId="16" pageNumber="261">et al.</emphasis>
(2009)
</bibRefCitation>
.
</paragraph>
<paragraph id="8BC536C91926FFEDFF0DFB44FE74FF5D" blockId="16.[132,775,217,1774]" pageId="16" pageNumber="261">
At the Miocene-Pliocene transition we see the appearance of the last currently admitted genera:
<taxonomicName id="4C7A4D4A1926FFEDFE79FB24FDFEFEE2" authorityName="Oyama" authorityYear="1964" box="[495,575,1238,1263]" class="Gastropoda" family="Bursidae" genus="Bursina" kingdom="Animalia" order="Littorinimorpha" pageId="16" pageNumber="265" phylum="Mollusca" rank="genus">
<emphasis id="B90EEADB1926FFEDFE79FB24FDFEFEE2" box="[495,575,1238,1263]" italics="true" pageId="16" pageNumber="261">Bursina</emphasis>
</taxonomicName>
in the Indo-Pacific and
<emphasis id="B90EEADB1926FFEDFF20FB04FECCFF1D" box="[182,269,1270,1296]" italics="true" pageId="16" pageNumber="261">Crossata</emphasis>
in western America (there is an unconfirmed occurrence of
<emphasis id="B90EEADB1926FFEDFEB4FAE4FEB8FF3D" box="[290,377,1302,1328]" italics="true" pageId="16" pageNumber="261">Crossata</emphasis>
in the middle Miocene (
<bibRefCitation id="EFEB4B381926FFEDFD02FAE4FCC1FF3D" author="BEU A. G." box="[660,768,1302,1328]" pageId="16" pageNumber="261" pagination="1 - 550" refId="ref10501" refString="BEU A. G. 2010. - Neogene tonnoidean gastropods of tropical and South America; a contribution to the Dominican Republic and Panama Paleontology projects and uplift of the Central American Isthmus. Bulletins of American Paleontology 377 - 378: 1 - 550." type="journal article" year="2010">Beu 2010</bibRefCitation>
;
<bibRefCitation id="EFEB4B381926FFEDFF12FAC7FE65FF42" author="POWELL II C. L. &amp; BERSCHAUER D. P." box="[132,420,1333,1359]" pageId="16" pageNumber="261" pagination="179 - 198" refId="ref12606" refString="POWELL II C. L. &amp; BERSCHAUER D. P. 2017. - Crossata (Gastropoda: Bursidae) in the eastern Pacific: a morphologic and paleontologic perspective. The Festivus 49 (3): 179 - 198." type="journal article" year="2017">Powell &amp; Berschauer 2017</bibRefCitation>
).
</paragraph>
<paragraph id="8BC536C91926FFEDFF0DFAA7FD28FCE3" blockId="16.[132,775,217,1774]" pageId="16" pageNumber="261">
Following the Messinian salinity crisis in the Mediterranean Sea, we observe the return of
<taxonomicName id="4C7A4D4A1926FFEDFD9FFA84FD79FF82" authorityName="" baseAuthorityName="Linnaeus" baseAuthorityYear="1758" box="[521,696,1397,1423]" class="Gastropoda" family="Bursidae" genus="Bursa" kingdom="Animalia" order="Littorinimorpha" pageId="16" pageNumber="261" phylum="Mollusca" rank="species" species="scrobilator">
<emphasis id="B90EEADB1926FFEDFD9FFA84FD79FF82" box="[521,696,1397,1423]" italics="true" pageId="16" pageNumber="261">Bursa scrobilator</emphasis>
</taxonomicName>
to this domain. In the Recent fauna it is the only bursid species in the Mediterranean.
<taxonomicName id="4C7A4D4A1926FFEDFECAFA44FDC0FFC2" baseAuthorityName="Perry" baseAuthorityYear="1811" box="[348,513,1462,1487]" class="Gastropoda" family="Bursidae" genus="Bursa" kingdom="Animalia" order="Littorinimorpha" pageId="16" pageNumber="261" phylum="Mollusca" rank="species" species="corrugata">
<emphasis id="B90EEADB1926FFEDFECAFA44FDC0FFC2" box="[348,513,1462,1487]" italics="true" pageId="16" pageNumber="261">Bursa corrugata</emphasis>
</taxonomicName>
returned briefly as well, but disappeared from the Mediterranean during the Pleistocene (
<bibRefCitation id="EFEB4B381926FFEDFF4DFA07FE64FC02" author="LANDAU B. &amp; HARZHAUSER M. &amp; BEU A. G." box="[219,421,1525,1551]" pageId="16" pageNumber="261" pagination="61 - 109" refId="ref11711" refString="LANDAU B., HARZHAUSER M. &amp; BEU A. G. 2009. - A Revision of the Tonnoidea (Caenogastropoda, Gastropoda) from the Miocene Paratethys and their palaeobiogeographic implications. Jahrbuch Der Geologischen Bundesanstalt 149: 61 - 109." type="journal article" year="2009">
Landau
<emphasis id="B90EEADB1926FFEDFEA5FA07FEA6FC02" box="[307,359,1525,1551]" italics="true" pageId="16" pageNumber="261">et al.</emphasis>
2009
</bibRefCitation>
). It was also during the Pliocene that
<taxonomicName id="4C7A4D4A1926FFEDFF23F9E7FEF4FC23" baseAuthorityName="Perry" baseAuthorityYear="1811" box="[181,309,1557,1582]" class="Gastropoda" family="Bursidae" genus="Bursa" kingdom="Animalia" order="Littorinimorpha" pageId="16" pageNumber="261" phylum="Mollusca" rank="species" species="corrugata">
<emphasis id="B90EEADB1926FFEDFF23F9E7FEF4FC23" box="[181,309,1557,1582]" italics="true" pageId="16" pageNumber="261">B. corrugata</emphasis>
</taxonomicName>
entered the western Atlantic, from which it reached the Eastern Pacific during the Pleistocene (
<bibRefCitation id="EFEB4B381926FFEDFD04F9C7FD36FC42" author="BEU A. G." box="[658,759,1589,1615]" pageId="16" pageNumber="261" pagination="1 - 550" refId="ref10501" refString="BEU A. G. 2010. - Neogene tonnoidean gastropods of tropical and South America; a contribution to the Dominican Republic and Panama Paleontology projects and uplift of the Central American Isthmus. Bulletins of American Paleontology 377 - 378: 1 - 550." type="journal article" year="2010">Beu 2010</bibRefCitation>
).
<taxonomicName id="4C7A4D4A1926FFEDFF12F9A7FEEAFC63" baseAuthorityName="Gmelin" baseAuthorityYear="1791" box="[132,299,1621,1647]" class="Gastropoda" family="Bursidae" genus="Aspa" kingdom="Animalia" order="Littorinimorpha" pageId="16" pageNumber="261" phylum="Mollusca" rank="species" species="marginata">
<emphasis id="B90EEADB1926FFEDFF12F9A7FEEAFC63" box="[132,299,1621,1647]" italics="true" pageId="16" pageNumber="261">Aspa marginata</emphasis>
</taxonomicName>
reappeared briefly in the Mediterranean at the beginning of the Pliocene and reached its maximum geographic extension during the late Pliocene/early Pleistocene by entering the Western Atlantic. At present it occurs only in the Eastern Atlantic and at some mid-Atlantic islands.
</paragraph>
</subSubSection>
</treatment>
</document>