167 lines
22 KiB
XML
167 lines
22 KiB
XML
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<mods:title id="D877CF469231B917F474088A0FB815A6">Fossil butterflies, calibration points and the molecular clock (Lepidoptera: Papilionoidea)</mods:title>
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<mods:namePart id="1E41DA46889DA236D792840CB7ED4EB2">Jong, Rienk De</mods:namePart>
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<mods:title id="E61463015AA01EEE6BDF831E70DC8B5E">Zootaxa</mods:title>
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<mods:date id="3D6F9656585ACA0BD6E91379BD67E2F5">2017</mods:date>
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<treatment id="03AA87D32858FFCFF7F0FF1FFE23B535" ID-DOI="http://doi.org/10.5281/zenodo.6046942" ID-GBIF-Taxon="130844882" ID-Zenodo-Dep="6046942" LSID="urn:lsid:plazi:treatment:03AA87D32858FFCFF7F0FF1FFE23B535" httpUri="http://treatment.plazi.org/id/03AA87D32858FFCFF7F0FF1FFE23B535" lastPageNumber="25" pageId="24" pageNumber="25">
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<subSubSection id="C319654E2858FFCFF7F0FF1FFF59B0C9" pageId="24" pageNumber="25" type="nomenclature">
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<paragraph id="8BBC36C52858FFCFF7F0FF1FFD48B08C" blockId="24.[151,747,152,178]" box="[151,747,152,178]" pageId="24" pageNumber="25">
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<heading id="D0F481A92858FFCFF7F0FF1FFD48B08C" bold="true" box="[151,747,152,178]" fontSize="11" level="1" pageId="24" pageNumber="25" reason="1">
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<emphasis id="B977EAD72858FFCFF7F0FF1FFD48B08C" bold="true" box="[151,747,152,178]" pageId="24" pageNumber="25">
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<emphasis id="B977EAD72858FFCFF7F0FF1FFF5CB08C" bold="true" box="[151,255,152,178]" italics="true" pageId="24" pageNumber="25">
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<taxonomicName id="4C034D462858FFCFF7F0FF1FFF58B08C" ID-CoL="5VQKD" box="[151,251,152,178]" class="Insecta" family="Pieridae" genus="Aporia" kingdom="Animalia" order="Lepidoptera" pageId="24" pageNumber="25" phylum="Arthropoda" rank="species" species="crataegi">crataegi</taxonomicName>
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.
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</emphasis>
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<taxonomicName id="4C034D462858FFCFF660FF1FFD48B08C" authority="Kernbach, 1967" authorityName="Kernbach" authorityYear="1967" box="[263,747,152,178]" class="Insecta" family="Pieridae" genus="Aporia" kingdom="Animalia" order="Lepidoptera" pageId="24" pageNumber="25" phylum="Arthropoda" rank="subSpecies" species="crataegi" subSpecies="fossilis">
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<emphasis id="B977EAD72858FFCFF660FF1FFDBEB08C" bold="true" box="[263,541,152,178]" italics="true" pageId="24" pageNumber="25">Aporia crataegi fossilis</emphasis>
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<bibRefCitation id="EF924B342858FFCFF543FF1FFD48B08C" author="Kernbach" box="[548,747,152,178]" pageId="24" pageNumber="25" refString="Kernbach, K. (1967) Uber die bisher im Pliozan von Willershausen gefundenen Schmetterlings- und Raupenreste. Berichte Naturhistorisches Gesellschaft Hannover, 111, 103 - 108." type="journal article" year="1967">Kernbach, 1967</bibRefCitation>
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</taxonomicName>
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</emphasis>
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||
</heading>
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</paragraph>
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<paragraph id="8BBC36C52858FFCFF7F0FF59FF59B0C9" blockId="24.[151,1437,222,1291]" box="[151,250,222,247]" pageId="24" pageNumber="25">
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<taxonomicName id="4C034D462858FFCFF7F0FF59FF54B0C9" box="[151,247,222,247]" class="Insecta" family="Pieridae" kingdom="Animalia" order="Lepidoptera" pageId="24" pageNumber="25" phylum="Arthropoda" rank="family">Pieridae</taxonomicName>
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.
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</paragraph>
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</subSubSection>
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<subSubSection id="C319654E2858FFCFF7A0FE84FE23B535" pageId="24" pageNumber="25" type="description">
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<paragraph id="8BBC36C52858FFCFF7A0FE84FC5AB125" blockId="24.[151,1437,222,1291]" box="[199,1017,258,283]" pageId="24" pageNumber="25">
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<materialsCitation id="3B6B3C982858FFCFF7A0FE84FC5AB125" ID-GBIF-Occurrence="1503739221" box="[199,1017,258,283]" country="Germany" location="Hesse" pageId="24" pageNumber="25" specimenCount="1" stateProvince="Hesse">
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<collectingCountry id="F31476552858FFCFF7A0FE84FE93B125" box="[199,304,259,283]" name="Germany" pageId="24" pageNumber="25">Germany</collectingCountry>
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,
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<collectingRegion id="49C7F8272858FFCFF65CFE84FE22B124" box="[315,385,259,282]" country="Germany" name="Hessen" pageId="24" pageNumber="25">Hesse</collectingRegion>
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,
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<collectingRegion id="49C7F8272858FFCFF6EBFE85FD82B125" box="[396,545,258,283]" country="Germany" name="Brandenburg" pageId="24" pageNumber="25">Brandenburg</collectingRegion>
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, Willershausen; Plazencian, late Pliocene.
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</materialsCitation>
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</paragraph>
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<paragraph id="8BBC36C52858FFCFF7A0FEA1FCACB101" blockId="24.[151,1437,222,1291]" box="[199,783,294,319]" pageId="24" pageNumber="25">Depository: GPUG (one specimen, 596-12[13589]).</paragraph>
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<paragraph id="8BBC36C52858FFCFF7A0FECDFBCCB1B9" blockId="24.[151,1437,222,1291]" pageId="24" pageNumber="25">
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Published figures:
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<bibRefCitation id="EF924B342858FFCFF6C5FECDFD54B15D" author="Branscheid" box="[418,759,330,355]" pageId="24" pageNumber="25" refString="Branscheid, F. (1968) Ein weiterer Schmetterlingsflugel aus dem Pliozan von Willershausen. Beihefte Berichte Naturhistorisches Gesellschaft Hannover, 6, 41 - 42." type="journal article" year="1968">
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Branscheid (1968:
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<figureCitation id="13382A402858FFCFF5E6FECDFD68B15D" box="[641,715,330,355]" captionStart="FIGURE 1" captionStartId="4.[151,250,1051,1073]" captionTargetBox="[167,1414,420,1017]" captionTargetId="figure@4.[157,1429,411,1026]" captionTargetPageId="4" captionText="FIGURE 1. Localities of fossil butterflies plotted ion a modern map. See Table 1 for further, complementary information." httpUri="https://zenodo.org/record/583184/files/figure.png" pageId="24" pageNumber="25">Figs 1</figureCitation>
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,
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<figureCitation id="13382A402858FFCFF5B9FECCFD4DB15D" box="[734,750,331,355]" captionStart="FIGURE 2" captionStartId="7.[151,250,1650,1672]" captionTargetBox="[175,1415,420,1615]" captionTargetId="figure@7.[151,1436,409,1627]" captionTargetPageId="7" captionText="FIGURE 2. Apomorphies, their placement on a tree, and the assignment of calibration points. See text for explanation." httpUri="https://zenodo.org/record/583185/files/figure.png" pageId="24" pageNumber="25">2</figureCitation>
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)
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</bibRefCitation>
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;
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<bibRefCitation id="EF924B342858FFCFF46EFECDFBF9B15D" author="Branscheid" box="[777,1114,330,355]" pageId="24" pageNumber="25" refString="Branscheid, F. (1969) Funde von Tagfaltern (Rhopalocera, Lepidopt.) im Pliozan von Willershausen. Berichte Naturhistorisches Gesellschaft Hannover, 113, 101 - 106." type="journal article" year="1969">
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Branscheid (1969:
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<figureCitation id="13382A402858FFCFF480FECDFBF2B15D" box="[999,1105,330,355]" captionStart-0="FIGURE 2" captionStart-1="FIGURES 3 – 4" captionStart-2="FIGURE 5" captionStartId-0="7.[151,250,1650,1672]" captionStartId-1="10.[151,264,1801,1823]" captionStartId-2="16.[151,250,1784,1806]" captionTargetBox-0="[175,1415,420,1615]" captionTargetBox-1="[165,1429,840,1774]" captionTargetBox-2="[238,1332,193,1743]" captionTargetId-0="figure@7.[151,1436,409,1627]" captionTargetId-1="figure@10.[154,1438,828,1788]" captionTargetId-2="figure@16.[238,1332,193,1754]" captionTargetPageId-0="7" captionTargetPageId-1="10" captionTargetPageId-2="16" captionText-0="FIGURE 2. Apomorphies, their placement on a tree, and the assignment of calibration points. See text for explanation." captionText-1="FIGURES 3 – 4. Wing venation of (A) Pareronia Bingham, 1907 (Pieridae), and (B) Hasora Moore, 1881 (Hesperiidae). See text for explanation." captionText-2="FIGURE 5. Ƒanessa amerindica Miller & Brown, 1989, holotype, collections of Invertebrate Paleontology, University of Florida. Photo by Roger Portell. Fig. 6. Pseudoneorina coulleti Nel & Descimon, 1984. France, Dép. Alpes-de-Haute- Provence, Dauphin, specimen in coll. M. Henrotay; photo by courtesy of M. Henrotay. Fig. 7. Archaeolycorea ferreirai Martins-Neto, 1989. Fig. 4 A of original description. Fig. 8. Oligodonta florissantensis Brown, 1976, holotype, collections of Invertebrate Paleontology, University of Florida. Photo by courtesy of Akito Kawahara. Fig. 9. Neorinella garciae Martins- Neto, Kucera-Santos, de Moraes Vieira & de Campos Fragoso, 1993. Fig. 3 A of original description. Fig. 10. Protocoeliades kristenseni de Jong, 2016, holotype, Zoologisk Museum, Copenhagen, Denmark. Photo by Department of Paleobiology, National Museum of Natural History, Washington DC, USA." httpUri-0="https://zenodo.org/record/583185/files/figure.png" httpUri-1="https://zenodo.org/record/583186/files/figure.png" httpUri-2="https://zenodo.org/record/583187/files/figure.png" pageId="24" pageNumber="25">Figs 2–5</figureCitation>
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)
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</bibRefCitation>
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;
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<bibRefCitation id="EF924B342858FFCFF30DFECDFA3BB15D" author="Kernbach" box="[1130,1432,330,355]" pageId="24" pageNumber="25" refString="Kernbach, K. (1967) Uber die bisher im Pliozan von Willershausen gefundenen Schmetterlings- und Raupenreste. Berichte Naturhistorisches Gesellschaft Hannover, 111, 103 - 108." type="journal article" year="1967">
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Kernbach (1967:
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<figureCitation id="13382A402858FFCFF250FECDFA2CB15D" box="[1335,1423,330,355]" captionStart="FIGURES 11 – 13" captionStartId="34.[151,264,1787,1809]" captionTargetBox="[211,1372,202,1743]" captionTargetId="figure@34.[174,1418,193,1750]" captionTargetPageId="34" captionText="FIGURES 11 – 13. Ƒanessa pluto Heer, 1849. Figures copied from (1875), Plate II, figures 1, 17 and 7, respectively. (A), copied by Scudder from original description by Heer; (B) and (C), figures sent to Scudder by Mr. Brunner de Wattenwyl, made after the same object. Note differences in wing shape and other details. Figs 14 – 16. Thaites ruminiana Scudder, 1875. Figures copied from Scudder (1875), Plate III, figures 9, 1 and 3, respectively. (A), underside of fossil; (B), reconstruction of venation; (C), reconstruction of markings. In (A) two separate subcostal veins have been drawn in the forewing, an impossible configuration in Lepidoptera, and apparently a mistake, corrected in (B) and (C). Fig. 17. Cyllo sepulta Boisduval, 1841, wing venation after Scudder (1875). Fig. 18. Butterfly fossil recorded by Wangrin (1939), but shown to be a fraud by Ansorge (2015). Photo kindly provided by J. Ansorge, Horst, Germany (see text for explanation)." httpUri="https://zenodo.org/record/583188/files/figure.png" pageId="24" pageNumber="25">Fig. 12</figureCitation>
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)
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</bibRefCitation>
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. Numerous figures of the extant species in identification guides for European butterflies.
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</paragraph>
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<paragraph id="8BBC36C52858FFCFF7A0FE15FB37B2F5" blockId="24.[151,1437,222,1291]" pageId="24" pageNumber="25">
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<bibRefCitation id="EF924B342858FFCFF7A0FE15FE2BB195" author="Kernbach" box="[199,392,402,427]" pageId="24" pageNumber="25" refString="Kernbach, K. (1967) Uber die bisher im Pliozan von Willershausen gefundenen Schmetterlings- und Raupenreste. Berichte Naturhistorisches Gesellschaft Hannover, 111, 103 - 108." type="journal article" year="1967">Kernbach (1967)</bibRefCitation>
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recorded one forewing and two hindwings from the clay of Willershausen as belonging to this extant Palaearctic species. The fossils are not described apart from the observation that the dark stripe between veins 1A+2A and CuA
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<quantity id="4CFB9B202858FFCFF6FBFE5CFE64B1CD" box="[412,455,474,499]" metricMagnitude="-2" metricUnit="m" metricValue="5.08" pageId="24" pageNumber="25" unit="in" value="2.0">2 in</quantity>
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the hindwing is visible, and that the specimens were apparently larger in the Pliocene. In the accompanying photo of the hindwing the venation is rather similar to that of
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<taxonomicName id="4C034D462858FFCFF321FE78FAE4B229" authority="Linnaeus" authorityName="Linnaeus" box="[1094,1351,510,535]" class="Insecta" family="Pieridae" genus="Aporia" kingdom="Animalia" order="Lepidoptera" pageId="24" pageNumber="25" phylum="Arthropoda" rank="species" species="crataegi">
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<emphasis id="B977EAD72858FFCFF321FE78FB61B228" box="[1094,1218,511,534]" italics="true" pageId="24" pageNumber="25">A. crataegi</emphasis>
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(Linnaeus)
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</taxonomicName>
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, except for Sc+R1 being much less strongly curved and the cell being distinctly shorter than in the recent species. Actually it is not clear what is typical about the
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<taxonomicName id="4C034D462858FFCFF509FDC0FD1FB260" box="[622,700,583,606]" class="Insecta" family="Pieridae" genus="Aporia" kingdom="Animalia" order="Lepidoptera" pageId="24" pageNumber="25" phylum="Arthropoda" rank="genus">
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<emphasis id="B977EAD72858FFCFF509FDC0FD1FB260" box="[622,700,583,606]" italics="true" pageId="24" pageNumber="25">Aporia</emphasis>
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</taxonomicName>
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hindwing venation, a similar venation also being found in the widespread recent genus
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<taxonomicName id="4C034D462858FFCFF6D3FDEBFE54B2BD" box="[436,503,620,643]" class="Insecta" family="Pieridae" genus="Pieris" kingdom="Animalia" order="Lepidoptera" pageId="24" pageNumber="25" phylum="Arthropoda" rank="genus">
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<emphasis id="B977EAD72858FFCFF6D3FDEBFE54B2BD" box="[436,503,620,643]" italics="true" pageId="24" pageNumber="25">Pieris</emphasis>
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</taxonomicName>
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and some other pierid genera. Also the stripe mentioned, which actually indicates the lost CuP, is not restricted to
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<taxonomicName id="4C034D462858FFCFF555FD08FD23B298" box="[562,640,655,678]" class="Insecta" family="Pieridae" genus="Aporia" kingdom="Animalia" order="Lepidoptera" pageId="24" pageNumber="25" phylum="Arthropoda" rank="genus">
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<emphasis id="B977EAD72858FFCFF555FD08FD23B298" box="[562,640,655,678]" italics="true" pageId="24" pageNumber="25">Aporia</emphasis>
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</taxonomicName>
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. Kernbach described the fossil as
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<taxonomicName id="4C034D462858FFCFF349FD08FAE6B299" box="[1070,1349,654,679]" class="Insecta" family="Pieridae" genus="Aporia" kingdom="Animalia" order="Lepidoptera" pageId="24" pageNumber="25" phylum="Arthropoda" rank="subSpecies" species="crataegi" subSpecies="fossilis">
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<emphasis id="B977EAD72858FFCFF349FD08FAE6B299" box="[1070,1349,654,679]" italics="true" pageId="24" pageNumber="25">Aporia crataegi fossilis</emphasis>
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</taxonomicName>
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, rather prematurely since it is not even certain that the fossil is conspecific with the extant species.
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</paragraph>
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<paragraph id="8BBC36C52858FFCFF7A0FD51FC3DB39D" blockId="24.[151,1437,222,1291]" pageId="24" pageNumber="25">
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||
Apparently, the forewing, mentioned but not described by
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||
<bibRefCitation id="EF924B342858FFCFF4E4FD51FBE5B2D1" author="Kernbach" box="[899,1094,726,751]" pageId="24" pageNumber="25" refString="Kernbach, K. (1967) Uber die bisher im Pliozan von Willershausen gefundenen Schmetterlings- und Raupenreste. Berichte Naturhistorisches Gesellschaft Hannover, 111, 103 - 108." type="journal article" year="1967">Kernbach (1967)</bibRefCitation>
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||
was reported by
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<bibRefCitation id="EF924B342858FFCFF279FD51FF46B32D" author="Branscheid" pageId="24" pageNumber="25" refString="Branscheid, F. (1968) Ein weiterer Schmetterlingsflugel aus dem Pliozan von Willershausen. Beihefte Berichte Naturhistorisches Gesellschaft Hannover, 6, 41 - 42." type="journal article" year="1968">Branscheid (1968)</bibRefCitation>
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. It shows the pierid apomorphy of M1 branching off the common stem of R4 and R5 (R3 is considered being lost by various authors). Sc is reconstructed as ending on the costa just beyond the end of the cell, but this may be a misinterpretation, even in fresh
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<taxonomicName id="4C034D462858FFCFF50DFCC4FD1BB364" box="[618,696,835,858]" class="Insecta" family="Pieridae" genus="Aporia" kingdom="Animalia" order="Lepidoptera" pageId="24" pageNumber="25" phylum="Arthropoda" rank="genus">
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<emphasis id="B977EAD72858FFCFF50DFCC4FD1BB364" box="[618,696,835,858]" italics="true" pageId="24" pageNumber="25">Aporia</emphasis>
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</taxonomicName>
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specimens it may be difficult to see where exactly Sc ends on the costa, this vein at its distal end running close to R1. Since the wing is incomplete, the general shape cannot be made out and its identification as belonging to
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<taxonomicName id="4C034D462858FFCFF506FC0CFD0CB39C" box="[609,687,907,930]" class="Insecta" family="Pieridae" genus="Aporia" kingdom="Animalia" order="Lepidoptera" pageId="24" pageNumber="25" phylum="Arthropoda" rank="genus">
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<emphasis id="B977EAD72858FFCFF506FC0CFD0CB39C" box="[609,687,907,930]" italics="true" pageId="24" pageNumber="25">Aporia</emphasis>
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</taxonomicName>
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is not without doubt.
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</paragraph>
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<paragraph id="8BBC36C52858FFCFF7A0FC29FE42B445" blockId="24.[151,1437,222,1291]" pageId="24" pageNumber="25">
|
||
Further wings from the same locality were described by
|
||
<bibRefCitation id="EF924B342858FFCFF439FC29FB92B3F9" author="Branscheid" box="[862,1073,942,967]" pageId="24" pageNumber="25" refString="Branscheid, F. (1969) Funde von Tagfaltern (Rhopalocera, Lepidopt.) im Pliozan von Willershausen. Berichte Naturhistorisches Gesellschaft Hannover, 113, 101 - 106." type="journal article" year="1969">Branscheid (1969)</bibRefCitation>
|
||
as most probably belonging to
|
||
<taxonomicName id="4C034D462858FFCFF7F0FC54FF46B3D4" box="[151,229,979,1002]" class="Insecta" family="Pieridae" genus="Aporia" kingdom="Animalia" order="Lepidoptera" pageId="24" pageNumber="25" phylum="Arthropoda" rank="genus">
|
||
<emphasis id="B977EAD72858FFCFF7F0FC54FF46B3D4" box="[151,229,979,1002]" italics="true" pageId="24" pageNumber="25">Aporia</emphasis>
|
||
</taxonomicName>
|
||
, although he was not sure about the (number of) species, since he found marked differences in size. For calibration purposes these fossils are hardly of interest.
|
||
<collectingRegion id="49C7F8272858FFCFF442FC70FCECB430" box="[805,847,1015,1038]" country="Cameroon" name="North" pageId="24" pageNumber="25">Not</collectingRegion>
|
||
only are they rather young, but since they cannot reliably be identified even to the subfamily level, we could only use them as calibration point at the root of the
|
||
<taxonomicName id="4C034D462858FFCFF7F0FBB9FF54B469" box="[151,247,1086,1111]" class="Insecta" family="Pieridae" kingdom="Animalia" order="Lepidoptera" pageId="24" pageNumber="25" phylum="Arthropoda" rank="family">Pieridae</taxonomicName>
|
||
. (For a recent study of the phylogeny, see
|
||
<bibRefCitation id="EF924B342858FFCFF5B5FBB9FC1CB469" author="Wahlberg" box="[722,959,1086,1111]" pageId="24" pageNumber="25" refString="Wahlberg, N., Rota, J., Braby, M. F. & Pierce, N. (2014) Revised systematics and higher classification of pierid butterflies (Lepidoptera: Pieridae) based on molecular data. Zoologica Scripta, 43, 641 - 650. https: // doi. org / 10.1111 / zsc. 12075" type="journal article" year="2014">
|
||
Wahlberg
|
||
<emphasis id="B977EAD72858FFCFF422FBB8FCD5B469" box="[837,886,1086,1111]" italics="true" pageId="24" pageNumber="25">et al</emphasis>
|
||
. 2014
|
||
</bibRefCitation>
|
||
, in which the late Pliocene appears far too recent for a divergence time.)
|
||
</paragraph>
|
||
<paragraph id="8BBC36C52858FFCFF7A0FB00FE23B535" blockId="24.[151,1437,222,1291]" pageId="24" pageNumber="25">
|
||
In view of the extreme scarcity of butterfly fossils it is remarkable that so many specimens of a single genus or maybe even species have been found in the same locality. A possible explanation could be gregarious behavior on wet spots involving so called mud puddling that can be observed in many butterflies, including
|
||
<taxonomicName id="4C034D462858FFCFF3DBFB48FACEB4D8" box="[1212,1389,1231,1254]" class="Insecta" family="Pieridae" genus="Aporia" kingdom="Animalia" order="Lepidoptera" pageId="24" pageNumber="25" phylum="Arthropoda" rank="species" species="crataegi">
|
||
<emphasis id="B977EAD72858FFCFF3DBFB48FACEB4D8" box="[1212,1389,1231,1254]" italics="true" pageId="24" pageNumber="25">Aporia crataegi</emphasis>
|
||
</taxonomicName>
|
||
and many other
|
||
<taxonomicName id="4C034D462858FFCFF67AFB75FEDEB535" box="[285,381,1266,1291]" class="Insecta" family="Pieridae" kingdom="Animalia" order="Lepidoptera" pageId="24" pageNumber="25" phylum="Arthropoda" rank="family">Pieridae</taxonomicName>
|
||
.
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |