316 lines
42 KiB
XML
316 lines
42 KiB
XML
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<mods:title id="4F22C289ABCE04D794EE6C61DF28CC5D">A molecular phylogeny of Speonemadus Jeannel, 1922 with description of two new species from Morocco (Coleoptera: Leiodidae: Cholevinae: Anemadini)</mods:title>
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<mods:namePart id="96AD7C60FBA1F9673EC1481805A555E9">Fresneda, Javier</mods:namePart>
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<mods:namePart id="47B81F57FB2337813AD65970A6236260">Faille, Arnaud</mods:namePart>
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<mods:namePart id="8B3B2B1665EF6AAEF03D3162E420FC65">Fery, Hans</mods:namePart>
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<mods:namePart id="3881631347656228B872861708E8C3BC">Ribera, Ignacio</mods:namePart>
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<treatment id="03FB8C32F803FFAFFF60FB84FF4D48D1" ID-DOI="http://doi.org/10.5281/zenodo.5927160" ID-GBIF-Taxon="154529609" ID-Zenodo-Dep="5927160" LSID="urn:lsid:plazi:treatment:03FB8C32F803FFAFFF60FB84FF4D48D1" httpUri="http://treatment.plazi.org/id/03FB8C32F803FFAFFF60FB84FF4D48D1" lastPageId="27" lastPageNumber="28" pageId="26" pageNumber="27">
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<subSubSection id="C3486EAFF803FFAEFF60FB84FDBE4F2F" box="[151,531,1052,1078]" pageId="26" pageNumber="27" type="nomenclature">
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<paragraph id="8BED3D24F803FFAEFF60FB84FDBE4F2F" blockId="26.[151,531,1052,1078]" box="[151,531,1052,1078]" pageId="26" pageNumber="27">
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<emphasis id="B926E136F803FFAEFF60FB84FDBE4F2F" bold="true" box="[151,531,1052,1078]" pageId="26" pageNumber="27">
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Biogeography of
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<taxonomicName id="4C5246A7F803FFAEFE9BFB84FDBE4F2F" ID-CoL="8X7VV" authorityName="Jeannel" authorityYear="1922" box="[364,531,1052,1078]" class="Insecta" family="Leiodidae" genus="Speonemadus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="genus">
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<emphasis id="B926E136F803FFAEFE9BFB84FDBE4F2F" bold="true" box="[364,531,1052,1078]" italics="true" pageId="26" pageNumber="27">Speonemadus</emphasis>
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</taxonomicName>
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</emphasis>
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</heading>
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</paragraph>
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<subSubSection id="C3486EAFF803FFAFFF60FBFAFF4D48D1" lastPageId="27" lastPageNumber="28" pageId="26" pageNumber="27" type="description">
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<paragraph id="8BED3D24F803FFAEFF60FBFAFCF34E6E" blockId="26.[151,1437,1122,2011]" pageId="26" pageNumber="27">
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The estimated phylogeny of
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<taxonomicName id="4C5246A7F803FFAEFE14FBFAFDD04F62" authorityName="Jeannel" authorityYear="1922" box="[483,637,1122,1147]" class="Insecta" family="Leiodidae" genus="Speonemadus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="genus">
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<emphasis id="B926E136F803FFAEFE14FBFAFDD04F62" box="[483,637,1122,1147]" italics="true" pageId="26" pageNumber="27">Speonemadus</emphasis>
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</taxonomicName>
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and the current distribution of the species are in agreement with the palaeogeography of the Betic-Rifean area during the Miocene (
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<figureCitation id="136921A1F803FFAEFC99FB1EFC594F86" box="[878,1012,1158,1183]" captionStart-0="FIGURES 40–41" captionStart-1="FIGURES 42–43" captionStartId-0="23.[151,264,1683,1705]" captionStartId-1="24.[151,264,1210,1232]" captionTargetBox-0="[428,1155,673,1661]" captionTargetBox-1="[426,1171,202,1189]" captionTargetId-0="figure@23.[409,1177,661,1662]" captionTargetId-1="figure@24.[416,1171,193,1189]" captionTargetPageId-0="23" captionTargetPageId-1="24" captionText-0="FIGURES 40–41. Palaeomaps with the reconstructed evolution of the distribution of Anemadus and Speonemadus at: (40) 18 Ma, Burdigalian; (41) 15 Ma, Langhian; (after Jolivet et al. 2006, modified)." captionText-1="FIGURES 42–43. Palaeomaps with the reconstructed evolution of the distribution of Anemadus and Speonemadus at: (42) 10 Ma, Late Serravallian-Tortonian; (43) 6–5.33 Ma Late Messinian, (after Jolivet et al. 2006, modified)." httpUri-0="https://zenodo.org/record/2617643/files/figure.png" httpUri-1="https://zenodo.org/record/2617645/files/figure.png" pageId="26" pageNumber="27">Figs 40–43</figureCitation>
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), as happens with other subterranean groups in the area (e.g. the ground beetle
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<taxonomicName id="4C5246A7F803FFAEFD8FFB32FCB14FDA" authorityName="Dejean" authorityYear="1831" box="[632,796,1194,1219]" class="Insecta" family="Carabidae" genus="Trechus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="species" species="fulvus">
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<emphasis id="B926E136F803FFAEFD8FFB32FCB14FDA" box="[632,796,1194,1219]" italics="true" pageId="26" pageNumber="27">Trechus fulvus</emphasis>
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</taxonomicName>
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-group,
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<bibRefCitation id="EFC340D5F803FFAEFC81FB32FB924FDA" author="Faille, A. & Andujar, C. & Fadrique, F. & Ribera, I." box="[886,1087,1194,1219]" pageId="26" pageNumber="27" pagination="1979 - 1990" refId="ref12562" refString="Faille, A., Andujar, C., Fadrique, F. & Ribera, I. (2014) Late Miocene origin of an Ibero-Maghrebian clade of ground beetles with multiple colonizations of the subterranean environment. Journal of Biogeography, 41, 1979 - 1990. https: // doi. org / 10.1111 / jbi. 12349" type="journal article" year="2014">
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Faille
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<emphasis id="B926E136F803FFAEFC48FB34FC5F4FDA" box="[959,1010,1194,1219]" italics="true" pageId="26" pageNumber="27">et al</emphasis>
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. 2014
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</bibRefCitation>
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). The geographic origin of the genus
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<taxonomicName id="4C5246A7F803FFAEFF13FB56FED34FFE" authorityName="Jeannel" authorityYear="1922" box="[228,382,1230,1255]" class="Insecta" family="Leiodidae" genus="Speonemadus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="genus">
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<emphasis id="B926E136F803FFAEFF13FB56FED34FFE" box="[228,382,1230,1255]" italics="true" pageId="26" pageNumber="27">Speonemadus</emphasis>
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</taxonomicName>
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cannot be unambiguously reconstructed, but considering the distribution of the presumed sister genus
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<taxonomicName id="4C5246A7F803FFAEFED3FB6AFE344E12" authorityName="Reitter" authorityYear="1885" box="[292,409,1266,1291]" class="Insecta" family="Leiodidae" genus="Anemadus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="genus">
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<emphasis id="B926E136F803FFAEFED3FB6AFE344E12" box="[292,409,1266,1291]" italics="true" pageId="26" pageNumber="27">Anemadus</emphasis>
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</taxonomicName>
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(Palaearctic, mostly from
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<collectingCountry id="F3457DB4F803FFAEFD3DFB6AFD504E12" box="[714,765,1266,1291]" name="Italy" pageId="26" pageNumber="27">Italy</collectingCountry>
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to the eastern Mediterranean and with species until
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<collectingCountry id="F3457DB4F803FFAEFAA2FB6BFA354E13" box="[1365,1432,1267,1290]" name="Japan" pageId="26" pageNumber="27">Japan</collectingCountry>
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; see
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<bibRefCitation id="EFC340D5F803FFAEFF36FA8FFEF44E36" author="Perreau, M." box="[193,345,1303,1327]" pageId="26" pageNumber="27" pagination="1 - 460" refId="ref14397" refString="Perreau, M. (2000) Catalogue des coleopteres Leiodidae Cholevinae et Platypsyllinae. Memoires de la Societe Entomologique de France, 4, 1 - 460." type="journal article" year="2000">Perreau 2000</bibRefCitation>
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), the origin is likely the western Palaearctic. Most Mediterranean islands are populated by species of
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<taxonomicName id="4C5246A7F803FFAEFF4CFAA2FE9D4E4A" authorityName="Reitter" authorityYear="1885" box="[187,304,1338,1363]" class="Insecta" family="Leiodidae" genus="Anemadus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="genus">
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<emphasis id="B926E136F803FFAEFF4CFAA2FE9D4E4A" box="[187,304,1338,1363]" italics="true" pageId="26" pageNumber="27">Anemadus</emphasis>
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</taxonomicName>
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(
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<collectingCountry id="F3457DB4F803FFAEFEB3FAA2FE334E4A" box="[324,414,1338,1363]" name="France" pageId="26" pageNumber="27">Corsica</collectingCountry>
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, Sardinia,
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<collectingCountry id="F3457DB4F803FFAEFDD6FAA3FDDF4E4A" box="[545,626,1339,1363]" name="Cyprus" pageId="26" pageNumber="27">Cyprus</collectingCountry>
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and some Greek Islands), which may have a late Messinian origin although there is no available phylogenetic data for the species.
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</paragraph>
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<paragraph id="8BED3D24F803FFAEFF30FA1BFB8F4C3F" blockId="26.[151,1437,1122,2011]" pageId="26" pageNumber="27">
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We estimated a crown age of
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<taxonomicName id="4C5246A7F803FFAEFDE7FA1AFD074E82" authorityName="Jeannel" authorityYear="1922" box="[528,682,1410,1435]" class="Insecta" family="Leiodidae" genus="Speonemadus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="genus">
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<emphasis id="B926E136F803FFAEFDE7FA1AFD074E82" box="[528,682,1410,1435]" italics="true" pageId="26" pageNumber="27">Speonemadus</emphasis>
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</taxonomicName>
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of ca. 10–11 Ma, with a split from the sampled
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<taxonomicName id="4C5246A7F803FFAEFB34FA1AFA954E82" authorityName="Reitter" authorityYear="1885" box="[1219,1336,1410,1435]" class="Insecta" family="Leiodidae" genus="Anemadus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="genus">
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<emphasis id="B926E136F803FFAEFB34FA1AFA954E82" box="[1219,1336,1410,1435]" italics="true" pageId="26" pageNumber="27">Anemadus</emphasis>
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</taxonomicName>
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at ca. 15 Ma (
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<figureCitation id="136921A1F803FFAEFF3CFA3EFEE24EA6" box="[203,335,1446,1471]" captionStart="FIGURES 40–41" captionStartId="23.[151,264,1683,1705]" captionTargetBox="[428,1155,673,1661]" captionTargetId="figure@23.[409,1177,661,1662]" captionTargetPageId="23" captionText="FIGURES 40–41. Palaeomaps with the reconstructed evolution of the distribution of Anemadus and Speonemadus at: (40) 18 Ma, Burdigalian; (41) 15 Ma, Langhian; (after Jolivet et al. 2006, modified)." httpUri="https://zenodo.org/record/2617643/files/figure.png" pageId="26" pageNumber="27">Figs 40–41</figureCitation>
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). This excludes the possibility of a colonisation of North Africa from Europe through the tectonic rafting of the Kabylian plates, which started to drift from the Iberian Peninsula at ca. 33 Ma, keeping some contact with the plates of the Balearic and Alboran islands until 26 Ma (
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<bibRefCitation id="EFC340D5F803FFAEFC9EFA76FBDC4D1E" author="Rosenbaum, G. & Lister, G. S. & Duboz, C." box="[873,1137,1518,1543]" pageId="26" pageNumber="27" pagination="107 - 130" refId="ref14784" refString="Rosenbaum, G., Lister, G. S. & Duboz, C. (2002) Reconstruction of the tectonic evolution of the western Mediterranean since the Oligocene. Journal of the Firtual Explorer, 8, 107 - 130. https: // doi. org / 10.3809 / jvirtex. 2002.00053" type="journal article" year="2002">
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Rosenbaum
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<emphasis id="B926E136F803FFAEFC01FA77FB8B4D1E" box="[1014,1062,1518,1543]" italics="true" pageId="26" pageNumber="27">et al</emphasis>
|
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. 2002
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</bibRefCitation>
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||
;
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<bibRefCitation id="EFC340D5F803FFAEFB8BFA76FA3C4D1E" author="Schettino, A. & Turco, E." box="[1148,1425,1518,1543]" pageId="26" pageNumber="27" pagination="1398 - 1423" refId="ref15121" refString="Schettino, A. & Turco, E. (2006) Plate kinematics of the Western Mediterranean region during the Oligocene and Early Miocene. Geophysical Journal International, 166, 1398 - 1423. https: // doi. org / 10.1111 / j. 1365 - 246 X. 2006.02997. x" type="journal article" year="2006">Schettino & Turco 2006</bibRefCitation>
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). Assuming a northern Mediterranean (i.e. European) origin of the genus, the separation between the ancestor of
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<taxonomicName id="4C5246A7F803FFAEFF60F9AEFE9C4D56" authorityName="Jeannel" authorityYear="1922" box="[151,305,1590,1615]" class="Insecta" family="Leiodidae" genus="Speonemadus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="genus">
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<emphasis id="B926E136F803FFAEFF60F9AEFE9C4D56" box="[151,305,1590,1615]" italics="true" pageId="26" pageNumber="27">Speonemadus</emphasis>
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</taxonomicName>
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and
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<taxonomicName id="4C5246A7F803FFAEFE9DF9AEFE724D56" authorityName="Reitter" authorityYear="1885" box="[362,479,1590,1615]" class="Insecta" family="Leiodidae" genus="Anemadus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="genus">
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<emphasis id="B926E136F803FFAEFE9DF9AEFE724D56" box="[362,479,1590,1615]" italics="true" pageId="26" pageNumber="27">Anemadus</emphasis>
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</taxonomicName>
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could have been mediated by the isolating effect of the Pyrenean chain (which was already formed by the late Oligocene, at ca. 23 Ma,
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<bibRefCitation id="EFC340D5F803FFAEFCF4F9C3FCDE4D6A" author="ISC" box="[771,883,1627,1651]" pageId="26" pageNumber="27" refId="ref13759" refString="ISC (2017) International Chronostratigraphic Chart. Fersion 2017 / 02. International Commission on Stratigraphy. Available from: http: // www. stratigraphy. org (accessed 9 May 2018)" type="url" year="2017">ISC 2017</bibRefCitation>
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) or, more likely, by the opening of the Rhone- Provençal basin (the "Basin miocène rhodano-provençal", BMRP, of
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<bibRefCitation id="EFC340D5F803FFAEFC68F9E7FB994D8E" author="Besson, D." box="[927,1076,1662,1687]" pageId="26" pageNumber="27" refId="ref11824" refString="Besson, D. (2005) Architecture du bassin rhodano-provencal miocene (Alpes, SE France): relations entre deformation, physiographie et sedimentation dans un bassin molassique d'avant-pays. Paris, Ecole Nationale Superieure des Mines de Paris, 438 pp." type="book" year="2005">Besson 2005</bibRefCitation>
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). The BMRP bordered the north side of the Alps from Marseille to
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<collectingRegion id="4996F3C6F803FFAEFDEAF93AFDC14DA2" box="[541,620,1698,1723]" country="Austria" name="Wien" pageId="26" pageNumber="27">Vienna</collectingRegion>
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(
|
||
<bibRefCitation id="EFC340D5F803FFAEFD8BF93AFC744DA2" author="Meulenkamp, J. E. & Sissingh, W." box="[636,985,1698,1723]" pageId="26" pageNumber="27" pagination="209 - 228" refId="ref14095" refString="Meulenkamp, J. E. & Sissingh, W. (2003) Tertiary palaeogeography and tectonostratigraphic evolution of the Northern and Southern Peri-Tethys platforms and the intermediate domains of the African-Eurasian convergent plate boundary zone. Palaeogeography, Palaeoclimatology, Palaeoecology, 196, 209 - 228. https: // doi. org / 10.1016 / S 0031 - 0182 (03) 00319 - 5" type="journal article" year="2003">Meulenkamp & Sissingh 2003</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFC340D5F803FFAEFC13F93BFBD44DA2" author="Besson, D." box="[996,1145,1698,1723]" pageId="26" pageNumber="27" refId="ref11824" refString="Besson, D. (2005) Architecture du bassin rhodano-provencal miocene (Alpes, SE France): relations entre deformation, physiographie et sedimentation dans un bassin molassique d'avant-pays. Paris, Ecole Nationale Superieure des Mines de Paris, 438 pp." type="book" year="2005">Besson 2005</bibRefCitation>
|
||
), with the stronger fluvial erosion dating from the Burdigalian (
|
||
<figureCitation id="136921A1F803FFAEFDCEF95EFD214DC6" box="[569,652,1734,1759]" captionStart="FIGURES 40–41" captionStartId="23.[151,264,1683,1705]" captionTargetBox="[428,1155,673,1661]" captionTargetId="figure@23.[409,1177,661,1662]" captionTargetPageId="23" captionText="FIGURES 40–41. Palaeomaps with the reconstructed evolution of the distribution of Anemadus and Speonemadus at: (40) 18 Ma, Burdigalian; (41) 15 Ma, Langhian; (after Jolivet et al. 2006, modified)." httpUri="https://zenodo.org/record/2617643/files/figure.png" pageId="26" pageNumber="27">Fig. 40</figureCitation>
|
||
), and a marine transgression inundating the region from the Langhian (
|
||
<figureCitation id="136921A1F803FFAEFF68F972FF544C1A" box="[159,249,1770,1795]" captionStart="FIGURES 40–41" captionStartId="23.[151,264,1683,1705]" captionTargetBox="[428,1155,673,1661]" captionTargetId="figure@23.[409,1177,661,1662]" captionTargetPageId="23" captionText="FIGURES 40–41. Palaeomaps with the reconstructed evolution of the distribution of Anemadus and Speonemadus at: (40) 18 Ma, Burdigalian; (41) 15 Ma, Langhian; (after Jolivet et al. 2006, modified)." httpUri="https://zenodo.org/record/2617643/files/figure.png" pageId="26" pageNumber="27">Fig. 41</figureCitation>
|
||
). This is in agreement with the hypotheseis of
|
||
<bibRefCitation id="EFC340D5F803FFAEFCCCF972FBDE4C1A" author="Giachino, P. M. & Vailati, D." box="[827,1139,1770,1795]" pageId="26" pageNumber="27" refId="ref13309" refString="Giachino, P. M. & Vailati, D. (1993) Revisione degli Anemadinae. Monografie di " Natura Bresciana " No. 18. Museo Civico di Scienze Naturali di Brescia, Brescia, 314 pp." type="book" year="1993">Giachino & Vailati (1993)</bibRefCitation>
|
||
of a separation between
|
||
<taxonomicName id="4C5246A7F803FFAEFF60F896FEA14C3E" authorityName="Reitter" authorityYear="1885" box="[151,268,1806,1831]" class="Insecta" family="Leiodidae" genus="Anemadus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="genus">
|
||
<emphasis id="B926E136F803FFAEFF60F896FEA14C3E" box="[151,268,1806,1831]" italics="true" pageId="26" pageNumber="27">Anemadus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C5246A7F803FFAEFEB5F896FE714C3E" authorityName="Jeannel" authorityYear="1922" box="[322,476,1806,1831]" class="Insecta" family="Leiodidae" genus="Speonemadus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="genus">
|
||
<emphasis id="B926E136F803FFAEFEB5F896FE714C3E" box="[322,476,1806,1831]" italics="true" pageId="26" pageNumber="27">Speonemadus</emphasis>
|
||
</taxonomicName>
|
||
during the late Miocene at both sides of the BMRP.
|
||
</paragraph>
|
||
<paragraph id="8BED3D24F803FFAFFF30F8AAFD0A4AED" blockId="26.[151,1437,1122,2011]" lastBlockId="27.[151,1436,151,968]" lastPageId="27" lastPageNumber="28" pageId="26" pageNumber="27">
|
||
The first cladogenesis separating the two basal clades of
|
||
<taxonomicName id="4C5246A7F803FFAEFC9AF8AAFBAA4C52" authorityName="Jeannel" authorityYear="1922" box="[877,1031,1842,1867]" class="Insecta" family="Leiodidae" genus="Speonemadus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="genus">
|
||
<emphasis id="B926E136F803FFAEFC9AF8AAFBAA4C52" box="[877,1031,1842,1867]" italics="true" pageId="26" pageNumber="27">Speonemadus</emphasis>
|
||
</taxonomicName>
|
||
(A and B) was estimated to have occurred in the Tortonian (ca. 10.5 Ma) (
|
||
<figureCitation id="136921A1F803FFAEFD96F8CEFD1B4C76" box="[609,694,1878,1903]" captionStart="FIGURES 42–43" captionStartId="24.[151,264,1210,1232]" captionTargetBox="[426,1171,202,1189]" captionTargetId="figure@24.[416,1171,193,1189]" captionTargetPageId="24" captionText="FIGURES 42–43. Palaeomaps with the reconstructed evolution of the distribution of Anemadus and Speonemadus at: (42) 10 Ma, Late Serravallian-Tortonian; (43) 6–5.33 Ma Late Messinian, (after Jolivet et al. 2006, modified)." httpUri="https://zenodo.org/record/2617645/files/figure.png" pageId="26" pageNumber="27">Fig. 42</figureCitation>
|
||
). Subsequently, in the early Messinian (ca. 6.5 Ma), a further split separated clades B1 and B2 (
|
||
<figureCitation id="136921A1F803FFAEFE16F8E2FD9A4C8A" box="[481,567,1914,1939]" captionStart="FIGURES 42–43" captionStartId="24.[151,264,1210,1232]" captionTargetBox="[426,1171,202,1189]" captionTargetId="figure@24.[416,1171,193,1189]" captionTargetPageId="24" captionText="FIGURES 42–43. Palaeomaps with the reconstructed evolution of the distribution of Anemadus and Speonemadus at: (42) 10 Ma, Late Serravallian-Tortonian; (43) 6–5.33 Ma Late Messinian, (after Jolivet et al. 2006, modified)." httpUri="https://zenodo.org/record/2617645/files/figure.png" pageId="26" pageNumber="27">Fig. 43</figureCitation>
|
||
). Each of these three clades has species both on the Iberian Peninsula and in North Africa. During the Tortonian and early Messinian the isolation of the highly dynamic Betic microplates has been hypothesised to have promoted the diversification of some groups (e.g.
|
||
<taxonomicName id="4C5246A7F803FFAEFC09F85AFB224CC2" authorityName="Thomson" authorityYear="1859" box="[1022,1167,1986,2011]" class="Insecta" family="Hydrochidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="family">Hydrochidae</taxonomicName>
|
||
water beetles, Hidalgo-
|
||
<bibRefCitation id="EFC340D5F802FFAFFF60FF0FFE0E4BA9" author="Hidalgo-Galiana, A. & Ribera, I." box="[151,419,151,176]" pageId="27" pageNumber="28" pagination="377 - 385" refId="ref13704" refString="Hidalgo-Galiana, A. & Ribera, I. (2011) Late Miocene diversification of the genus Hydrochus (Coleoptera, Hydrochidae) in the west Mediterranean area. Molecular Phylogenetics and Evolution, 59, 377 - 385. https: // doi. org / 10.1016 / j. ympev. 2011.01.018" type="journal article" year="2011">Galiana & Ribera 2011</bibRefCitation>
|
||
, or
|
||
<taxonomicName id="4C5246A7F802FFAFFE27FF0FFDCA4BA9" authorityName="Thomson" authorityYear="1875" box="[464,615,151,176]" class="Insecta" family="Carabidae" genus="Mesocarabus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="genus">
|
||
<emphasis id="B926E136F802FFAFFE27FF0FFDCA4BA9" box="[464,615,151,176]" italics="true" pageId="27" pageNumber="28">Mesocarabus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C5246A7F802FFAFFD56FF0FFD544BA9" box="[673,761,151,176]" class="Insecta" family="Carabidae" genus="Trechus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="genus">
|
||
<emphasis id="B926E136F802FFAFFD56FF0FFD544BA9" box="[673,761,151,176]" italics="true" pageId="27" pageNumber="28">Trechus</emphasis>
|
||
</taxonomicName>
|
||
ground beetles,
|
||
<bibRefCitation id="EFC340D5F802FFAFFC40FF0FFB3A4BA9" author="Andujar, C. & Gomez-Zurita, J. & Rasplus, J. - Y. & Serrano, J." box="[951,1175,151,176]" pageId="27" pageNumber="28" pagination="787 - 804" refId="ref11565" refString="Andujar, C., Gomez-Zurita, J., Rasplus, J. - Y. & Serrano, J. (2012) Molecular systematics and evolution of the subgenus Mesocarabus Thomson, 1875 (Coleoptera: Carabidae: Carabus), based on mitochondrial and nuclear DNA. Zoological Journal of the Linnean Society, 166, 787 - 804. https: // doi. org / 10.1111 / j. 1096 - 3642.2012.00866. x" type="journal article" year="2012">
|
||
Andújar
|
||
<emphasis id="B926E136F802FFAFFBEBFF01FBE24BA9" box="[1052,1103,151,176]" italics="true" pageId="27" pageNumber="28">et al</emphasis>
|
||
. 2012
|
||
</bibRefCitation>
|
||
and
|
||
<bibRefCitation id="EFC340D5F802FFAFFB26FF0FFA3A4BA9" author="Faille, A. & Andujar, C. & Fadrique, F. & Ribera, I." box="[1233,1431,151,176]" pageId="27" pageNumber="28" pagination="1979 - 1990" refId="ref12562" refString="Faille, A., Andujar, C., Fadrique, F. & Ribera, I. (2014) Late Miocene origin of an Ibero-Maghrebian clade of ground beetles with multiple colonizations of the subterranean environment. Journal of Biogeography, 41, 1979 - 1990. https: // doi. org / 10.1111 / jbi. 12349" type="journal article" year="2014">
|
||
Faille
|
||
<emphasis id="B926E136F802FFAFFAEDFF01FAE04BA9" box="[1306,1357,151,176]" italics="true" pageId="27" pageNumber="28">et al</emphasis>
|
||
. 2014
|
||
</bibRefCitation>
|
||
, respectively); this could also have been the case in
|
||
<taxonomicName id="4C5246A7F802FFAFFD16FF24FCD64BCC" authorityName="Jeannel" authorityYear="1922" box="[737,891,188,213]" class="Insecta" family="Leiodidae" genus="Speonemadus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="genus">
|
||
<emphasis id="B926E136F802FFAFFD16FF24FCD64BCC" box="[737,891,188,213]" italics="true" pageId="27" pageNumber="28">Speonemadus</emphasis>
|
||
</taxonomicName>
|
||
, with the isolation of three lineages (A, B1 and B2) either in
|
||
<collectingCountry id="F3457DB4F802FFAFFEDDFF47FEC34BE1" box="[298,366,223,248]" name="Georgia" pageId="27" pageNumber="28">Iberia</collectingCountry>
|
||
, North Africa or some of the Betic-Rifean plates. During the salinity crisis of the late Messinian all these areas became united, likely allowing the expansion of the species through the Strait of
|
||
<collectingCountry id="F3457DB4F802FFAFFB20FE9CFA934A04" box="[1239,1342,260,285]" name="Gibraltar" pageId="27" pageNumber="28">Gibraltar</collectingCountry>
|
||
. The reopening of the
|
||
<collectingCountry id="F3457DB4F802FFAFFEB4FEBFFE054A59" box="[323,424,295,320]" name="Gibraltar" pageId="27" pageNumber="28">Gibraltar</collectingCountry>
|
||
strait at the end of the Messinian salinity crisis 5.33 Ma (
|
||
<bibRefCitation id="EFC340D5F802FFAFFBCDFEBFFA384A59" author="Garcia-Castellanos, D. & Estrada, F. & Jimenez-Munt, I. & Gorini, C. & Fernandez, M. & Verges, J. & Vicente, R. de" box="[1082,1429,295,320]" pageId="27" pageNumber="28" pagination="778 - 781" refId="ref13189" refString="Garcia-Castellanos, D., Estrada, F., Jimenez-Munt, I., Gorini, C., Fernandez, M., Verges, J. & Vicente, R. de (2009) Catastrophic flood of the Mediterranean after the Messinian salinity crisis. Nature, 462, 778 - 781. https: // doi. org / 10.1038 / nature 08555" type="journal article" year="2009">
|
||
García-Castellanos
|
||
<emphasis id="B926E136F802FFAFFAEEFEB1FAE64A59" box="[1305,1355,295,320]" italics="true" pageId="27" pageNumber="28">et al</emphasis>
|
||
. 2009
|
||
</bibRefCitation>
|
||
) isolated the Iberian from the North African populations of these three widespread ancestral
|
||
<taxonomicName id="4C5246A7F802FFAFFB65FED4FA814A7C" authorityName="Jeannel" authorityYear="1922" box="[1170,1324,332,357]" class="Insecta" family="Leiodidae" genus="Speonemadus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="genus">
|
||
<emphasis id="B926E136F802FFAFFB65FED4FA814A7C" box="[1170,1324,332,357]" italics="true" pageId="27" pageNumber="28">Speonemadus</emphasis>
|
||
</taxonomicName>
|
||
, resulting in an almost simultaneous split into three clades: (1) clade A, the Iberian
|
||
<taxonomicName id="4C5246A7F802FFAFFC1DFEE8FBD74A91" baseAuthorityName="Perris" baseAuthorityYear="1864" box="[1002,1146,367,392]" class="Insecta" family="Leiodidae" genus="Speonemadus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="species" species="clathratus">
|
||
<emphasis id="B926E136F802FFAFFC1DFEE8FBD74A91" box="[1002,1146,367,392]" italics="true" pageId="27" pageNumber="28">S. clathratus</emphasis>
|
||
</taxonomicName>
|
||
and the species of the
|
||
<taxonomicName id="4C5246A7F802FFAFFA70FEE8FF534AB4" baseAuthorityName="Uhagon" baseAuthorityYear="1898" class="Insecta" family="Leiodidae" genus="Speonemadus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="species" species="escalerai">
|
||
<emphasis id="B926E136F802FFAFFA70FEE8FF534AB4" italics="true" pageId="27" pageNumber="28">S. escalerai</emphasis>
|
||
</taxonomicName>
|
||
-group vs. the North African
|
||
<taxonomicName id="4C5246A7F802FFAFFDB1FE0DFD154AB4" authority="Fresneda & Faille & Fery & Ribera, 2019" authorityName="Fresneda & Faille & Fery & Ribera" authorityYear="2019" box="[582,696,404,429]" class="Insecta" family="Leiodidae" genus="Speonemadus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="species" species="brusteli" status="sp. nov.">
|
||
<emphasis id="B926E136F802FFAFFDB1FE0DFD154AB4" box="[582,696,404,429]" italics="true" pageId="27" pageNumber="28">S. brusteli</emphasis>
|
||
</taxonomicName>
|
||
<emphasis id="B926E136F802FFAFFD37FE0CFCAD4AB4" bold="true" box="[704,768,404,429]" pageId="27" pageNumber="28">
|
||
<taxonomicNameLabel id="A2155C4DF802FFAFFD37FE0CFCAD4AB4" box="[704,768,404,429]" pageId="27" pageNumber="28" rank="species">sp. n.</taxonomicNameLabel>
|
||
</emphasis>
|
||
and
|
||
<taxonomicName id="4C5246A7F802FFAFFCC1FE0DFC764AB5" baseAuthorityName="Jeannel" baseAuthorityYear="1936" box="[822,987,405,428]" class="Insecta" family="Leiodidae" genus="Speonemadus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="species" species="maroccanus">
|
||
<emphasis id="B926E136F802FFAFFCC1FE0DFC764AB5" box="[822,987,405,428]" italics="true" pageId="27" pageNumber="28">S. maroccanus</emphasis>
|
||
</taxonomicName>
|
||
(plus most likely
|
||
<taxonomicName id="4C5246A7F802FFAFFB50FE0DFAFD4AB4" box="[1191,1360,404,429]" class="Insecta" family="Leiodidae" genus="Speonemadus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="species" species="orchesioides">
|
||
<emphasis id="B926E136F802FFAFFB50FE0DFAFD4AB4" box="[1191,1360,404,429]" italics="true" pageId="27" pageNumber="28">S. orchesioides</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C5246A7F802FFAFFA70FE0DFEB74AC9" baseAuthorityName="Reiche" baseAuthorityYear="1864" class="Insecta" family="Leiodidae" genus="Speonemadus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="species" species="subcostatus">
|
||
<emphasis id="B926E136F802FFAFFA70FE0DFEB74AC9" italics="true" pageId="27" pageNumber="28">S. subcostatus</emphasis>
|
||
</taxonomicName>
|
||
); (2) clade B1, the Iberian
|
||
<taxonomicName id="4C5246A7F802FFAFFDAEFE20FD604AC9" authorityName="Kraatz" authorityYear="1870" box="[601,717,439,464]" class="Insecta" family="Leiodidae" genus="Choleva" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="species" species="gracilis">
|
||
<emphasis id="B926E136F802FFAFFDAEFE20FD604AC9" box="[601,717,439,464]" italics="true" pageId="27" pageNumber="28">S. gracilis</emphasis>
|
||
</taxonomicName>
|
||
vs. the North African
|
||
<taxonomicName id="4C5246A7F802FFAFFC22FE20FBFC4AC9" authorityName="Peyerimhoff" authorityYear="1917" box="[981,1105,440,464]" class="Insecta" family="Leiodidae" genus="Anemadus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="species" species="tenuipes">
|
||
<emphasis id="B926E136F802FFAFFC22FE20FBFC4AC9" box="[981,1105,440,464]" italics="true" pageId="27" pageNumber="28">S. tenuipes</emphasis>
|
||
</taxonomicName>
|
||
; (3) clade B2, the Iberian
|
||
<taxonomicName id="4C5246A7F802FFAFFA70FE20FEB84AEC" authorityName="Heyden" authorityYear="1870" class="Insecta" family="Leiodidae" genus="Catops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="species" species="vandalitiae">
|
||
<emphasis id="B926E136F802FFAFFA70FE20FEB84AEC" italics="true" pageId="27" pageNumber="28">S. vandalitiae</emphasis>
|
||
</taxonomicName>
|
||
vs. the North African
|
||
<taxonomicName id="4C5246A7F802FFAFFDE6FE45FD0D4AEC" box="[529,672,476,501]" class="Insecta" family="Leiodidae" genus="Speonemadus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="species" species="pulchellus">
|
||
<emphasis id="B926E136F802FFAFFDE6FE45FD0D4AEC" box="[529,672,476,501]" italics="true" pageId="27" pageNumber="28">S. pulchellus</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</paragraph>
|
||
<paragraph id="8BED3D24F802FFAFFF30FE67FDF849D4" blockId="27.[151,1436,151,968]" pageId="27" pageNumber="28">
|
||
This scenario would require the colonisation of south
|
||
<collectingCountry id="F3457DB4F802FFAFFCDAFE67FCC34901" box="[813,878,511,536]" name="Georgia" pageId="27" pageNumber="28">Iberia</collectingCountry>
|
||
by
|
||
<taxonomicName id="4C5246A7F802FFAFFC6CFD98FBEC4901" baseAuthorityName="Jeannel" baseAuthorityYear="1936" box="[923,1089,512,536]" class="Insecta" family="Leiodidae" genus="Speonemadus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="species" species="maroccanus">
|
||
<emphasis id="B926E136F802FFAFFC6CFD98FBEC4901" box="[923,1089,512,536]" italics="true" pageId="27" pageNumber="28">S. maroccanus</emphasis>
|
||
</taxonomicName>
|
||
during the Pleistocene, and of Sicily—and possible continental Italy—during the Plio-Pleistocene. There is no information on the timing of these colonisations, they may have been related to the climatic fluctuations during the Pliocene, with first a temperature and precipitation higher than at present (
|
||
<bibRefCitation id="EFC340D5F802FFAFFD96FDF4FCE9499D" author="Dowsett, H. & Barron, J. A. & Poore, R. Z. & Thompson, R. S. & Cronin, T. M. & Ishman, S. E. & Willard, D. A." box="[609,836,620,645]" pageId="27" pageNumber="28" refId="ref12384" refString="Dowsett, H., Barron, J. A., Poore, R. Z., Thompson, R. S., Cronin, T. M., Ishman, S. E. & Willard, D. A. (1999) Middle Pliocene paleoenvironmental reconstruction: PRISM 2. U. S. Geological Survey Open File Report 99 - 535. https: // doi. org / 10.3133 / ofr 99535" type="book" year="1999">
|
||
Dowsett
|
||
<emphasis id="B926E136F802FFAFFD3FFDF5FCAC499C" box="[712,769,620,645]" italics="true" pageId="27" pageNumber="28">et al.</emphasis>
|
||
1999
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFC340D5F802FFAFFCA6FDF4FBED499D" author="Haywood, A. M. & Sellwood, B. W. & Valdes, P. J." box="[849,1088,620,645]" pageId="27" pageNumber="28" pagination="1063 - 1066" refId="ref13433" refString="Haywood, A. M., Sellwood, B. W. & Valdes, P. J. (2000) Regional warming: Pliocene (3 Ma) paleoclimate of Europe and the Mediterranean. Geology, 28, 1063 - 1066. https: // doi. org / 10.1130 / 0091 - 7613 (2000) 28 % 3 C 1063: RWPMPO % 3 E 2.0. CO; 2" type="journal article" year="2000">
|
||
Haywood
|
||
<emphasis id="B926E136F802FFAFFC32FDF5FC55499C" box="[965,1016,620,645]" italics="true" pageId="27" pageNumber="28">et al</emphasis>
|
||
. 2000
|
||
</bibRefCitation>
|
||
) and then the establishment of the Mediterranean climate at ca. 3.2 Ma (
|
||
<bibRefCitation id="EFC340D5F802FFAFFD85FD08FD4E49B1" author="Suc, J. - P." box="[626,739,656,680]" pageId="27" pageNumber="28" pagination="429 - 432" refId="ref15421" refString="Suc, J. - P. (1984) Origin and evolution of the Mediterranean vegetation and climate in Europe. Nature, 307, 429 - 432. https: // doi. org / 10.1038 / 307429 a 0" type="journal article" year="1984">Suc 1984</bibRefCitation>
|
||
), or they may have been facilitated by the descent of the sea level during the Pleistocene glaciations.
|
||
</paragraph>
|
||
<paragraph id="8BED3D24F802FFAFFF30FD4FFF4D48D1" blockId="27.[151,1436,151,968]" pageId="27" pageNumber="28">
|
||
Within the Iberian Peninsula, in the late Pliocene
|
||
<taxonomicName id="4C5246A7F802FFAFFD07FD40FCD349E9" baseAuthorityName="Perris" baseAuthorityYear="1864" box="[752,894,727,752]" class="Insecta" family="Leiodidae" genus="Speonemadus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="species" species="clathratus">
|
||
<emphasis id="B926E136F802FFAFFD07FD40FCD349E9" box="[752,894,727,752]" italics="true" pageId="27" pageNumber="28">S. clathratus</emphasis>
|
||
</taxonomicName>
|
||
may have been isolated from the ancestor of the
|
||
<taxonomicName id="4C5246A7F802FFAFFF60FD65FEB3480C" baseAuthorityName="Uhagon" baseAuthorityYear="1898" box="[151,286,764,789]" class="Insecta" family="Leiodidae" genus="Speonemadus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="species" species="escalerai">
|
||
<emphasis id="B926E136F802FFAFFF60FD65FEB3480C" box="[151,286,764,789]" italics="true" pageId="27" pageNumber="28">S. escalerai</emphasis>
|
||
</taxonomicName>
|
||
-group by the Gualdalquivir valley, either due to the onset of the Mediterranean climate or by the marine transgressions and inland seas that inundated most of the valley until the Middle Pleistocene (
|
||
<bibRefCitation id="EFC340D5F802FFAFFAE5FCB8FF794845" author="Torres, T. & Ortiz, J. E. & Soler, V. & Reyes, E. & Delgado, A. & Valle, M. & Llamas, J. F. & Cobo, R. & Julia, R. & Badal, E. & Garcia de la Morena, M. A. & Garcia-Martinez, M. J. & Fernandez-Gianotti, J. & Calvo, J. P. & Cortes, A." pageId="27" pageNumber="28" pagination="151 - 185" refId="ref15630" refString="Torres, T., Ortiz, J. E., Soler, V., Reyes, E., Delgado, A., Valle, M., Llamas, J. F., Cobo, R., Julia, R., Badal, E., Garcia de la Morena, M. A., Garcia-Martinez, M. J., Fernandez-Gianotti, J., Calvo, J. P. & Cortes, A. (2003) Pleistocene lacustrine basin of the east domain of guadix-baza basin (Granada, Spain): sedimentology, chronostratigraphy, and palaeoenvironment. In: Valero-Garces, B. L. (Ed.), Limnogeologia en Espana: un homenaje a Kerry R. Kelts. Consejo Superior de Investigaciones Cientificas, Madrid, pp. 151 - 185." type="book chapter" year="2003">
|
||
Torres
|
||
<emphasis id="B926E136F802FFAFFA94FCB9FA384821" box="[1379,1429,799,824]" italics="true" pageId="27" pageNumber="28">et al</emphasis>
|
||
. 2003
|
||
</bibRefCitation>
|
||
). The increased seasonality and aridification, as well as the Pleistocene glaciations and the associated climatic changes, may have limited the surface movement of some species of
|
||
<taxonomicName id="4C5246A7F802FFAFFC34FCFFFBF04899" authorityName="Jeannel" authorityYear="1922" box="[963,1117,871,896]" class="Insecta" family="Leiodidae" genus="Speonemadus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="genus">
|
||
<emphasis id="B926E136F802FFAFFC34FCFFFBF04899" box="[963,1117,871,896]" italics="true" pageId="27" pageNumber="28">Speonemadus</emphasis>
|
||
</taxonomicName>
|
||
, promoting the isolation of peripheral populations in the subterranean medium and giving origin to the current species of the
|
||
<taxonomicName id="4C5246A7F802FFAFFAFBFC15FA3E48BC" baseAuthorityName="Uhagon" baseAuthorityYear="1898" box="[1292,1427,908,933]" class="Insecta" family="Leiodidae" genus="Speonemadus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="species" species="escalerai">
|
||
<emphasis id="B926E136F802FFAFFAFBFC15FA3E48BC" box="[1292,1427,908,933]" italics="true" pageId="27" pageNumber="28">S. escalerai</emphasis>
|
||
</taxonomicName>
|
||
- group.
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |