456 lines
57 KiB
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456 lines
57 KiB
XML
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<mods:title id="5C7752B5872009F5DAE557E7209F2DC1">Fossil butterflies, calibration points and the molecular clock (Lepidoptera: Papilionoidea)</mods:title>
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<treatment id="03AA87D32852FFC4F7F0FDCFFDF1B5DA" ID-DOI="http://doi.org/10.5281/zenodo.6046920" ID-GBIF-Taxon="130844892" ID-Zenodo-Dep="6046920" LSID="urn:lsid:plazi:treatment:03AA87D32852FFC4F7F0FDCFFDF1B5DA" httpUri="http://treatment.plazi.org/id/03AA87D32852FFC4F7F0FDCFFDF1B5DA" lastPageId="19" lastPageNumber="20" pageId="18" pageNumber="19">
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<subSubSection id="C319654E2852FFC5F7F0FDCFFDC1B299" pageId="18" pageNumber="19" type="nomenclature">
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<paragraph id="8BBC36C52852FFC5F7F0FDCFFD02B25C" blockId="18.[151,673,584,610]" box="[151,673,584,610]" pageId="18" pageNumber="19">
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<heading id="D0F481A92852FFC5F7F0FDCFFD02B25C" bold="true" box="[151,673,584,610]" fontSize="11" level="1" pageId="18" pageNumber="19" reason="1">
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<emphasis id="B977EAD72852FFC5F7F0FDCFFD02B25C" bold="true" box="[151,673,584,610]" pageId="18" pageNumber="19">
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<emphasis id="B977EAD72852FFC5F7F0FDCFFEBAB25C" bold="true" box="[151,281,584,610]" italics="true" pageId="18" pageNumber="19">
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<taxonomicName id="4C034D462852FFC5F7F0FDCFFEB7B25C" box="[151,276,584,610]" class="Insecta" family="Papilionidae" genus="Doritites" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="species" species="bosniaskii">bosniaskii</taxonomicName>
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.
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</emphasis>
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<taxonomicName id="4C034D462852FFC5F647FDCFFD02B25C" authority="Rebel, 1898" authorityName="Rebel" authorityYear="1898" box="[288,673,584,610]" class="Insecta" family="Papilionidae" genus="Doritites" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="species" species="bosniaskii">
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<emphasis id="B977EAD72852FFC5F647FDCFFDAAB25C" bold="true" box="[288,521,584,610]" italics="true" pageId="18" pageNumber="19">Doritites bosniaskii</emphasis>
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<bibRefCitation id="EF924B342852FFC5F577FDCFFD02B25C" author="Rebel" box="[528,673,584,610]" pageId="18" pageNumber="19" refString="Rebel, H. (1898) Fossile Lepidopteren aus der Miocan-Formation von Gabbro. Sitzungsberichte der Akademie der Wissenschaften Wien, 107, 731 - 745." type="journal article" year="1898">Rebel, 1898</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="8BBC36C52852FFC5F7F0FD09FDC1B299" blockId="18.[151,1437,654,2011]" box="[151,610,654,679]" pageId="18" pageNumber="19">
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<taxonomicName id="4C034D462852FFC5F7F0FD09FE8BB299" box="[151,296,654,679]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="family">Papilionidae</taxonomicName>
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:
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<taxonomicName id="4C034D462852FFC5F655FD09FE60B299" box="[306,451,654,679]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="subFamily" subFamily="Parnassiinae">Parnassiinae</taxonomicName>
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:
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<taxonomicName id="4C034D462852FFC5F6ABFD09FDFDB299" box="[460,606,654,679]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="tribe" tribe="Luehdorfiini">Luehdorfiini</taxonomicName>
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.
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</paragraph>
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</subSubSection>
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<subSubSection id="C319654E2852FFC4F7A0FD35FDF1B5DA" lastPageId="19" lastPageNumber="20" pageId="18" pageNumber="19" type="description">
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<paragraph id="8BBC36C52852FFC5F7A0FD35FCE7B2F5" blockId="18.[151,1437,654,2011]" box="[199,836,690,715]" pageId="18" pageNumber="19">Italy, Gabbro (near Pisa), Italy; Messinian, late Miocene.</paragraph>
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<paragraph id="8BBC36C52852FFC5F7A0FD51FB10B2D1" blockId="18.[151,1437,654,2011]" box="[199,1203,726,751]" pageId="18" pageNumber="19">Depository: NHMW (holotype, 1898/0013/0001; counterpart holotype, 1898/0013/0002).</paragraph>
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<paragraph id="8BBC36C52852FFC5F7A0FD7DFD93B309" blockId="18.[151,1437,654,2011]" pageId="18" pageNumber="19">
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Published figures:
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<bibRefCitation id="EF924B342852FFC5F6FDFD7DFD0BB32D" author="Demoulin" box="[410,680,762,787]" pageId="18" pageNumber="19" refString="Demoulin, G. (1975) A propos de deux Papilionidae mediterraneens fossiles. Bulletin Cercle des Lepidopteristes de Belgique, 4, 96 - 101." type="journal article" year="1975">
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Demoulin (1975:
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<figureCitation id="13382A402852FFC5F505FD7DFD3FB32D" box="[610,668,762,787]" 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="18" pageNumber="19">Fig 1</figureCitation>
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)
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</bibRefCitation>
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;
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<bibRefCitation id="EF924B342852FFC5F5D2FD7CFC53B32D" author="Murata" box="[693,1008,762,787]" pageId="18" pageNumber="19" refString="Murata, Y. (1998) Notes on the evolutionary process of butterflies and butterfly fossils. Butterflies, 20 (part 1 & 2), 4 - 17 & 27 - 40. [in Japanese with English summary]" type="journal article" year="1998">Murata (1998: Figs. 35, 36)</bibRefCitation>
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;
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<bibRefCitation id="EF924B342852FFC5F49BFD7DFF40B309" author="Ponomarenko" pageId="18" pageNumber="19" refString="Ponomarenko, A. G. & Schultz, O. (1988) Kataloge der wissenschaftlichen Sammlung des Naturhistorischen Museums in Wien. Bd. 6. Palaozoologie. Heft 1. Natuurhistorisches Museum, Wien, 39 pp., 14 pls." type="book" year="1988">
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Ponomarenko & Schultz (1988: Pl. 7
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<figureCitation id="13382A402852FFC5F7F0FC99FF74B309" box="[151,215,798,823]" 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="18" pageNumber="19">Fig. 1</figureCitation>
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)
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</bibRefCitation>
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;
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<bibRefCitation id="EF924B342852FFC5F797FC99FD88B309" author="Rebel" box="[240,555,798,823]" pageId="18" pageNumber="19" refString="Rebel, H. (1898) Fossile Lepidopteren aus der Miocan-Formation von Gabbro. Sitzungsberichte der Akademie der Wissenschaften Wien, 107, 731 - 745." type="journal article" year="1898">
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Rebel (1898: Pl. I
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<figureCitation id="13382A402852FFC5F6D9FC99FD81B309" box="[446,546,798,823]" captionStart-0="FIGURE 1" captionStart-1="FIGURE 2" captionStart-2="FIGURES 3 – 4" captionStartId-0="4.[151,250,1051,1073]" captionStartId-1="7.[151,250,1650,1672]" captionStartId-2="10.[151,264,1801,1823]" captionTargetBox-0="[167,1414,420,1017]" captionTargetBox-1="[175,1415,420,1615]" captionTargetBox-2="[165,1429,840,1774]" captionTargetId-0="figure@4.[157,1429,411,1026]" captionTargetId-1="figure@7.[151,1436,409,1627]" captionTargetId-2="figure@10.[154,1438,828,1788]" captionTargetPageId-0="4" captionTargetPageId-1="7" captionTargetPageId-2="10" captionText-0="FIGURE 1. Localities of fossil butterflies plotted ion a modern map. See Table 1 for further, complementary information." captionText-1="FIGURE 2. Apomorphies, their placement on a tree, and the assignment of calibration points. See text for explanation." captionText-2="FIGURES 3 – 4. Wing venation of (A) Pareronia Bingham, 1907 (Pieridae), and (B) Hasora Moore, 1881 (Hesperiidae). See text for explanation." httpUri-0="https://zenodo.org/record/583184/files/figure.png" httpUri-1="https://zenodo.org/record/583185/files/figure.png" httpUri-2="https://zenodo.org/record/583186/files/figure.png" pageId="18" pageNumber="19">Figs 1–3</figureCitation>
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)
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</bibRefCitation>
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.
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</paragraph>
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<paragraph id="8BBC36C52852FFC5F7A0FCC5FBADB7AD" blockId="18.[151,1437,654,2011]" pageId="18" pageNumber="19">
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Reasonably well preserved butterfly: head, thorax, abdomen, right forewing partly overlapped by right hindwing, basal parts of left wings, all seen from the underside. Forewing radial formula: 1, 2, 3+(4+5). The posterior margin of the cell is reconstructed as quadrifid, a papilionid autapomorphy. A cubital spur (also a papilionid autapomorphy) is not apparent. Further apomorphies of the
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<taxonomicName id="4C034D462852FFC5F4C0FC29FB95B3F9" box="[935,1078,942,967]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="family">Papilionidae</taxonomicName>
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, such as the second anal vein on the forewing running to the dorsum, and the single anal vein on the hindwing, are not apparent in the fossil.
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<bibRefCitation id="EF924B342852FFC5F23CFC55FE83B431" author="Rebel" pageId="18" pageNumber="19" refString="Rebel, H. (1898) Fossile Lepidopteren aus der Miocan-Formation von Gabbro. Sitzungsberichte der Akademie der Wissenschaften Wien, 107, 731 - 745." type="journal article" year="1898">Rebel (1898: 736)</bibRefCitation>
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remarked: “Die länglichen Hinterflügel zeigen, wie in der Gattung
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<taxonomicName id="4C034D462852FFC5F32BFC70FAAFB430" authority="Latr." authorityName="Latr." box="[1100,1292,1015,1038]" class="Insecta" family="Papilionidae" genus="Parnassius" kingdom="Animalia" order="Lepidoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="genus">
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<emphasis id="B977EAD72852FFC5F32BFC70FB6BB430" box="[1100,1224,1015,1038]" italics="true" pageId="18" pageNumber="19">Parnassius</emphasis>
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Latr.
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</taxonomicName>
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, einen stark ausgeschnittenen Innenrandtheil [...]” ("As in the genus
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<taxonomicName id="4C034D462852FFC5F46DFB9BFC66B40C" authority="Latr." authorityName="Latr." box="[778,965,1051,1075]" class="Insecta" family="Papilionidae" genus="Parnassius" kingdom="Animalia" order="Lepidoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="genus">
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<emphasis id="B977EAD72852FFC5F46DFB9BFC25B40D" box="[778,902,1052,1075]" italics="true" pageId="18" pageNumber="19">Parnassius</emphasis>
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Latr.
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</taxonomicName>
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, the oblong hindwings have a dorsum that strongly curves inwards […]"), but the fossil is not decisive here (see also
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<bibRefCitation id="EF924B342852FFC5F4BFFBB8FBC9B469" author="Zeuner" box="[984,1130,1086,1111]" pageId="18" pageNumber="19" refString="Zeuner, F. E. (1961) Notes on the evolution of the Rhopalocera (Lep.). Ferhandlungen des XI. Internationalen Kongresses fur Entomologie, 1 (1960), 310 - 313." type="journal article" year="1961">Zeuner 1961</bibRefCitation>
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). At the tip of the abdomen a sphragis is visible, indicating that the fossil is a fertilized female (or at least that it has mated). A sphragis is found in members of the
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<taxonomicName id="4C034D462852FFC5F601FB01FE51B4A1" box="[358,498,1158,1183]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="subFamily" subFamily="Parnassiinae">Parnassiinae</taxonomicName>
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and some
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<taxonomicName id="4C034D462852FFC5F50EFB01FD5AB4A1" box="[617,761,1158,1183]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="subFamily" subFamily="Papilioninae">Papilioninae</taxonomicName>
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. It could, at most, be an apomorphy at the family level and is plesiomorphic within the subfamilies. Probably as a convergent development, a sphragis also can be found in some
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<taxonomicName id="4C034D462852FFC5F7F0FB49FE92B4D9" box="[151,305,1230,1255]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="family">Nymphalidae</taxonomicName>
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, such as the genus
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<taxonomicName id="4C034D462852FFC5F561FB48FDC0B4D8" box="[518,611,1231,1254]" class="Insecta" family="Nymphalidae" genus="Actinote" kingdom="Animalia" order="Lepidoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="genus">
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<emphasis id="B977EAD72852FFC5F561FB48FDC0B4D8" box="[518,611,1231,1254]" italics="true" pageId="18" pageNumber="19">Actinote</emphasis>
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</taxonomicName>
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(see
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<bibRefCitation id="EF924B342852FFC5F5F9FB49FCCEB4D9" author="Paluch" box="[670,877,1230,1255]" pageId="18" pageNumber="19" refString="Paluch, M., Casagrande, M. M. & Mielke, O. H. H. (2003) Tampao genital de Actinote Hubner, como carater toxonomico (Lepidoptera, Nymphalidae, Acraeinae). Revista Brasileira de Entomologia, 47, 573 - 580. https: // doi. org / 10.1590 / S 0085 - 56262003000400007" type="journal article" year="2003">
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Paluch
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<emphasis id="B977EAD72852FFC5F596FB48FC81B4D9" box="[753,802,1230,1255]" italics="true" pageId="18" pageNumber="19">et al</emphasis>
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. 2003
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</bibRefCitation>
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||
). With its rounded and untailed wings the fossil is similar to species of the genera
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<taxonomicName id="4C034D462852FFC5F54FFB73FD07B535" box="[552,676,1268,1291]" class="Insecta" family="Papilionidae" genus="Parnassius" kingdom="Animalia" order="Lepidoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="genus">
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<emphasis id="B977EAD72852FFC5F54FFB73FD07B535" box="[552,676,1268,1291]" italics="true" pageId="18" pageNumber="19">Parnassius</emphasis>
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</taxonomicName>
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and
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<taxonomicName id="4C034D462852FFC5F58DFB75FC98B535" box="[746,827,1266,1291]" class="Insecta" family="Papilionidae" genus="Archon" kingdom="Animalia" order="Lepidoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="genus">
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<emphasis id="B977EAD72852FFC5F58DFB75FC98B535" box="[746,827,1266,1291]" italics="true" pageId="18" pageNumber="19">Archon</emphasis>
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</taxonomicName>
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of the subfamily
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<taxonomicName id="4C034D462852FFC5F347FB75FB13B535" box="[1056,1200,1266,1291]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="subFamily" subFamily="Parnassiinae">Parnassiinae</taxonomicName>
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. Monophyly of the Parnassinae is only weakly based on characters that are not visible in the fossil (
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<bibRefCitation id="EF924B342852FFC5F32DFA91FA88B511" author="Ackery" box="[1098,1323,1302,1327]" pageId="18" pageNumber="19" refString="Ackery, P. R., de Jong, R. & Vane-Wright, R. I. (1999) The butterflies: Hedyloidea, Hesperioidea and Papilionoidea. In: Kristensen, N. P. (Ed.), Lepidoptera, Moths and Butterflies. 1. Evolution, Systematics, and Biogeography. Handbook of Zoology. 4 (35). Lepidoptera. De Gruyter, Berlin, pp. 263 - 300." type="book chapter" year="1999">
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Ackery
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<emphasis id="B977EAD72852FFC5F3CEFA90FB7EB511" box="[1193,1245,1302,1327]" italics="true" pageId="18" pageNumber="19">et al</emphasis>
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. 1999
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</bibRefCitation>
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). Yet, the similarity with species in this subfamily is so striking, that it most probably belongs here. On the basis of recent analyses based on mitochondrial DNA (
|
||
<bibRefCitation id="EF924B342852FFC5F515FAD8FCF0B549" author="Omoto" box="[626,851,1374,1399]" pageId="18" pageNumber="19" refString="Omoto, K., Katoh, T., Chichvarkhin, A. & Yagi, T. (2004) Molecular systematics and evolution of the " Apollo " butterflies of the genus Parnassius (Lepidoptera: Papilionidae), based on mitochondrial DNA sequence data. Gene, 326, 141 - 147. https: // doi. org / 10.1016 / j. gene. 2003.10.020" type="journal article" year="2004">
|
||
Omoto
|
||
<emphasis id="B977EAD72852FFC5F5A9FAD8FCA6B549" box="[718,773,1374,1399]" italics="true" pageId="18" pageNumber="19">et al</emphasis>
|
||
. 2004
|
||
</bibRefCitation>
|
||
) and mitochondrial and nuclear DNA as well as morphology (
|
||
<bibRefCitation id="EF924B342852FFC5F657FA05FDA0B5A5" author="Nazari" box="[304,515,1410,1435]" pageId="18" pageNumber="19" refString="Nazari, V., Zakharov, E. V. & Sperling, F. A. H. (2007) Phylogeny, historical biogeography, and taxonomic ranking of Parnassiinae (Lepidoptera: Papilionidae) based on morphology and seven genes. Molecular Phylogenetics and Evolution, 42, 131 - 156." type="journal article" year="2007">
|
||
Nazari
|
||
<emphasis id="B977EAD72852FFC5F6E2FA03FE14B5A5" box="[389,439,1410,1435]" italics="true" pageId="18" pageNumber="19">et al</emphasis>
|
||
. 2007
|
||
</bibRefCitation>
|
||
), three monophyletic groups can be recognized in the subfamily, that were given tribal rank by
|
||
<bibRefCitation id="EF924B342852FFC5F65BFA21FDBCB581" box="[316,543,1446,1471]" pageId="18" pageNumber="19" refString="Nazari, V., Zakharov, E. V. & Sperling, F. A. H. (2007) Phylogeny, historical biogeography, and taxonomic ranking of Parnassiinae (Lepidoptera: Papilionidae) based on morphology and seven genes. Molecular Phylogenetics and Evolution, 42, 131 - 156." type="journal article">
|
||
Nazari
|
||
<emphasis id="B977EAD72852FFC5F6F7FA20FE60B581" box="[400,451,1446,1471]" italics="true" pageId="18" pageNumber="19">et al</emphasis>
|
||
. (2007)
|
||
</bibRefCitation>
|
||
as follows:
|
||
<taxonomicName id="4C034D462852FFC5F5C8FA21FC8FB581" box="[687,812,1446,1471]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="tribe" tribe="Parnassiini">Parnassiini</taxonomicName>
|
||
+(
|
||
<taxonomicName id="4C034D462852FFC5F425FA21FC77B581" box="[834,980,1446,1471]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="tribe" tribe="Luehdorfiini">Luehdorfiini</taxonomicName>
|
||
+
|
||
<taxonomicName id="4C034D462852FFC5F484FA21FBC7B581" box="[995,1124,1446,1471]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="tribe" tribe="Zerynthiini">Zerynthiini</taxonomicName>
|
||
). In the fossil the precostal (=humeral) vein on the hindwing is simple. This is found in
|
||
<taxonomicName id="4C034D462852FFC5F459FA4DFC68B5DD" box="[830,971,1482,1507]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="tribe" tribe="Luehdorfiini">Luehdorfiini</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C034D462852FFC5F365FA4DFB20B5DD" box="[1026,1155,1482,1507]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="tribe" tribe="Zerynthiini">Zerynthiini</taxonomicName>
|
||
, while in
|
||
<taxonomicName id="4C034D462852FFC5F396FA4DFACFB5DD" box="[1265,1388,1482,1507]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="tribe" tribe="Parnassiini">Parnassiini</taxonomicName>
|
||
this vein is forked. The unforked condition is supposed to be apomorphic with respect to the forked condition by
|
||
<bibRefCitation id="EF924B342852FFC5F7F0F995FEF1B615" author="Hancock" box="[151,338,1554,1579]" pageId="18" pageNumber="19" refString="Hancock, D. L. (1983) Classification of the Papilionidae (Lepidoptera): a phylogenetic approach. Smithersia, 2, 1 - 48." type="journal article" year="1983">Hancock (1983)</bibRefCitation>
|
||
, who based his conclusions on morphological characters only. According to modern analyses based on DNA (
|
||
<bibRefCitation id="EF924B342852FFC5F63CF9B1FDF4B671" author="Zakharov" box="[347,599,1590,1615]" pageId="18" pageNumber="19" refString="Zakharov, E. V., Caterino, M. S. & Sperling, F. A. (2004) Molecular Phylogeny, Historical Biogeography, and Divergence Time Estimates for Swallowtail Butterflies of the Genus Papilio (Lepidoptera: Papilionidae). Systematic Biology, 53, 1 - 23. https: // doi. org / 10.1080 / 10635150490423403" type="journal article" year="2004">
|
||
Zakharov
|
||
<emphasis id="B977EAD72852FFC5F6B3F9B0FDAAB671" box="[468,521,1590,1615]" italics="true" pageId="18" pageNumber="19">et al</emphasis>
|
||
. 2004
|
||
</bibRefCitation>
|
||
) or on DNA and morphological characters (
|
||
<bibRefCitation id="EF924B342852FFC5F30FF9B1FAC6B671" author="Simonsen" box="[1128,1381,1590,1615]" pageId="18" pageNumber="19" refString="Simonsen, T. J., Zakharov, E. V., Djernaes, M., Cotton, A. M., Vane-Wright, R. I. & Sperling, F. A. H. (2011) Phylogenetics and divergence times of Papilionae (Lepidoptera) with special reference to the enigmatic genera Teinopalpus and Meandrusa. Cladistics, 27, 113 - 137." type="journal article" year="2011">
|
||
Simonsen
|
||
<emphasis id="B977EAD72852FFC5F383F9B0FAB8B671" box="[1252,1307,1590,1615]" italics="true" pageId="18" pageNumber="19">et al</emphasis>
|
||
. 2011
|
||
</bibRefCitation>
|
||
) the relationship of the subfamilies of the
|
||
<taxonomicName id="4C034D462852FFC5F523F9DDFD73B64D" box="[580,720,1626,1651]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="family">Papilionidae</taxonomicName>
|
||
is
|
||
<taxonomicName id="4C034D462852FFC5F592F9DDFCD1B64D" box="[757,882,1626,1651]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="subFamily" subFamily="Baroniinae">Baroniinae</taxonomicName>
|
||
+(
|
||
<taxonomicName id="4C034D462852FFC5F4EFF9DDFBB4B64D" box="[904,1047,1626,1651]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="subFamily" subFamily="Parnassiinae">Parnassiinae</taxonomicName>
|
||
+
|
||
<taxonomicName id="4C034D462852FFC5F341F9DDFB15B64D" box="[1062,1206,1626,1651]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="subFamily" subFamily="Papilioninae">Papilioninae</taxonomicName>
|
||
), although
|
||
<bibRefCitation id="EF924B342852FFC5F253F9DDFEAFB6A9" pageId="18" pageNumber="19" refString="Nazari, V., Zakharov, E. V. & Sperling, F. A. H. (2007) Phylogeny, historical biogeography, and taxonomic ranking of Parnassiinae (Lepidoptera: Papilionidae) based on morphology and seven genes. Molecular Phylogenetics and Evolution, 42, 131 - 156." type="journal article">
|
||
Nazari
|
||
<emphasis id="B977EAD72852FFC5F2EFF9DBFF0EB6A9" italics="true" pageId="18" pageNumber="19">et al</emphasis>
|
||
. (2007)
|
||
</bibRefCitation>
|
||
, who also used molecular as well as morphological data found a sister group relationship between
|
||
<taxonomicName id="4C034D462852FFC5F7F0F925FEB1B685" box="[151,274,1698,1723]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="subFamily" subFamily="Baroniinae">Baroniinae</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C034D462852FFC5F62EF925FE7AB685" box="[329,473,1698,1723]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="subFamily" subFamily="Parnassiinae">Parnassiinae</taxonomicName>
|
||
. A forked precostal vein is found in
|
||
<taxonomicName id="4C034D462852FFC5F411F925FC57B685" box="[886,1012,1698,1723]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="subFamily" subFamily="Baroniinae">Baroniinae</taxonomicName>
|
||
, part of the
|
||
<taxonomicName id="4C034D462852FFC5F31CF925FA3FB685" authority="(Parnassiini)" baseAuthorityName="Parnassiini" box="[1147,1436,1698,1723]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="subFamily" subFamily="Parnassiinae">Parnassiinae (Parnassiini)</taxonomicName>
|
||
and in most of the
|
||
<taxonomicName id="4C034D462852FFC5F60AF941FE5EB6E1" box="[365,509,1734,1759]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="subFamily" subFamily="Papilioninae">Papilioninae</taxonomicName>
|
||
. In
|
||
<taxonomicName id="4C034D462852FFC5F54FF941FD27B6E1" box="[552,644,1734,1759]" class="Insecta" family="Pieridae" kingdom="Animalia" order="Lepidoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="family">Pieridae</taxonomicName>
|
||
the condition is unforked, if present; in
|
||
<taxonomicName id="4C034D462852FFC5F328F941FB45B6E1" box="[1103,1254,1734,1759]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="family">Nymphalidae</taxonomicName>
|
||
both conditions occur, in
|
||
<taxonomicName id="4C034D462852FFC5F664F96DFEDDB73D" box="[259,382,1770,1795]" class="Insecta" family="Riodinidae" kingdom="Animalia" order="Lepidoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="family">Riodinidae</taxonomicName>
|
||
the precostal vein is unforked, in
|
||
<taxonomicName id="4C034D462852FFC5F46EF96DFC2AB73D" box="[777,905,1770,1795]" class="Insecta" family="Lycaenidae" kingdom="Animalia" order="Lepidoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="family">Lycaenidae</taxonomicName>
|
||
generally absent. Such a distribution makes it difficult to attach much value to this character. If the unforked condition is apomorphic within the
|
||
<taxonomicName id="4C034D462852FFC5F395F889FA22B719" box="[1266,1409,1806,1831]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="family">Papilionidae</taxonomicName>
|
||
, it evolved in
|
||
<taxonomicName id="4C034D462852FFC5F674F8B5FE2DB775" box="[275,398,1842,1867]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="tribe" tribe="Parnassiini">Parnassiini</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C034D462852FFC5F6A3F8B5FDF0B775" box="[452,595,1842,1867]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="subFamily" subFamily="Papilioninae">Papilioninae</taxonomicName>
|
||
, and it evolved several times in distantly related groups of
|
||
<taxonomicName id="4C034D462852FFC5F382F8B5FA23B775" box="[1253,1408,1842,1867]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="family">Nymphalidae</taxonomicName>
|
||
. If so, the simple condition of the precostal vein in
|
||
<taxonomicName id="4C034D462852FFC5F5C9F8D0FC2AB751" box="[686,905,1878,1903]" class="Insecta" family="Papilionidae" genus="Doritites" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="species" species="bosniaskii">
|
||
<emphasis id="B977EAD72852FFC5F5C9F8D0FC2AB751" box="[686,905,1878,1903]" italics="true" pageId="18" pageNumber="19">Doritites bosniaskii</emphasis>
|
||
</taxonomicName>
|
||
is a plesiomorphy and need not point to a close relationship with
|
||
<taxonomicName id="4C034D462852FFC5F63CF8FDFE7AB7AD" box="[347,473,1914,1939]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="tribe" tribe="Zerynthiini">Zerynthiini</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C034D462852FFC5F568F8FDFD03B7AD" box="[527,672,1914,1939]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="tribe" tribe="Luehdorfiini">Luehdorfiini</taxonomicName>
|
||
, but does not contradict it either.
|
||
</paragraph>
|
||
<paragraph id="8BBC36C52852FFC4F7A0F819FC54B203" blockId="18.[151,1437,654,2011]" lastBlockId="19.[151,1436,151,1509]" lastPageId="19" lastPageNumber="20" pageId="18" pageNumber="19">
|
||
The forewing has a dark outer border, which according to
|
||
<bibRefCitation id="EF924B342852FFC5F40EF819FB9AB789" author="Rebel" box="[873,1081,1950,1975]" pageId="18" pageNumber="19" refString="Rebel, H. (1898) Fossile Lepidopteren aus der Miocan-Formation von Gabbro. Sitzungsberichte der Akademie der Wissenschaften Wien, 107, 731 - 745." type="journal article" year="1898">Rebel (1898: 737)</bibRefCitation>
|
||
was probably hyaline, but this idea was apparently prompted by the assumption that the fossil was in the direct ancestry of
|
||
<taxonomicName id="4C034D462852FFC5F3DFF843FA97B7E5" box="[1208,1332,1988,2011]" class="Insecta" family="Papilionidae" genus="Parnassius" kingdom="Animalia" order="Lepidoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="genus">
|
||
<emphasis id="B977EAD72852FFC5F3DFF843FA97B7E5" box="[1208,1332,1988,2011]" italics="true" pageId="18" pageNumber="19">Parnassius</emphasis>
|
||
</taxonomicName>
|
||
. Further, there are four bands, one subapical between cell and apex and extending from costa to halfway between M1 and M2, one at the end of the cell, one across the central part of the cell, and one more basally in the cell. Elements of these markings can be found throughout the
|
||
<taxonomicName id="4C034D462853FFC4F5E1FF58FCB1B0C6" box="[646,786,223,248]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="subFamily" subFamily="Parnassiinae">Parnassiinae</taxonomicName>
|
||
and in some
|
||
<taxonomicName id="4C034D462853FFC4F4C1FF58FB91B0C6" box="[934,1074,223,248]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="subFamily" subFamily="Papilioninae">Papilioninae</taxonomicName>
|
||
as well (for instance,
|
||
<taxonomicName id="4C034D462853FFC4F241FF58FA37B0C6" box="[1318,1428,223,248]" class="Insecta" family="Papilionidae" genus="Iphiclides" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="genus">
|
||
<emphasis id="B977EAD72853FFC4F241FF58FA37B0C6" box="[1318,1428,223,248]" italics="true" pageId="19" pageNumber="20">Iphiclides</emphasis>
|
||
</taxonomicName>
|
||
, some
|
||
<taxonomicName id="4C034D462853FFC4F7BFFE83FE88B123" box="[216,299,260,285]" class="Insecta" family="Papilionidae" genus="Papilio" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="genus">
|
||
<emphasis id="B977EAD72853FFC4F7BFFE83FE88B123" box="[216,299,260,285]" italics="true" pageId="19" pageNumber="20">Papilio</emphasis>
|
||
</taxonomicName>
|
||
species), though the exact place may differ a little. It is considered to be part of the original papilionid design (
|
||
<bibRefCitation id="EF924B342853FFC4F794FEA0FE7DB17E" author="Schwanwitsch" box="[243,478,295,320]" pageId="19" pageNumber="20" refString="Schwanwitsch, B. N. (1943) Wing pattern in papilionid Lepidoptera. Entomologist, 76, 201 - 203." type="journal article" year="1943">Schwanwitsch 1943</bibRefCitation>
|
||
), and the fact that it resembles for instance the markings of
|
||
<taxonomicName id="4C034D462853FFC4F3D2FEAEFEB6B15A" authority="Eversmann" authorityName="Eversmann" class="Insecta" family="Papilionidae" genus="Parnassius" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="species" species="delphius">
|
||
<emphasis id="B977EAD72853FFC4F3D2FEAEFA3FB17E" box="[1205,1436,295,320]" italics="true" pageId="19" pageNumber="20">Parnassius delphius</emphasis>
|
||
Eversmann
|
||
</taxonomicName>
|
||
(cf.
|
||
<bibRefCitation id="EF924B342853FFC4F637FECBFE75B15A" author="Rebel" box="[336,470,332,357]" pageId="19" pageNumber="20" refString="Rebel, H. (1898) Fossile Lepidopteren aus der Miocan-Formation von Gabbro. Sitzungsberichte der Akademie der Wissenschaften Wien, 107, 731 - 745." type="journal article" year="1898">Rebel 1898</bibRefCitation>
|
||
) cannot be considered evidence of close relationship with the latter. The hindwing shows a dark outer border of which the proximal edge is less well defined than in the forewing, and a dark oblique band over the discocellular veins continuing to the costa. In
|
||
<taxonomicName id="4C034D462853FFC4F42DFE13FC75B193" box="[842,982,404,429]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="subFamily" subFamily="Parnassiinae">Parnassiinae</taxonomicName>
|
||
a similar oblique band is only found in
|
||
<taxonomicName id="4C034D462853FFC4F7F0FE30FEB0B1EE" box="[151,275,439,464]" class="Insecta" family="Papilionidae" genus="Luehdorfia" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="genus">
|
||
<emphasis id="B977EAD72853FFC4F7F0FE30FEB0B1EE" box="[151,275,439,464]" italics="true" pageId="19" pageNumber="20">Luehdorfia</emphasis>
|
||
</taxonomicName>
|
||
though elements of such a band can be found in
|
||
<taxonomicName id="4C034D462853FFC4F45EFE3FFC02B1F1" box="[825,929,440,463]" class="Insecta" family="Papilionidae" genus="Sericinus" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="genus">
|
||
<emphasis id="B977EAD72853FFC4F45EFE3FFC02B1F1" box="[825,929,440,463]" italics="true" pageId="19" pageNumber="20">Sericinus</emphasis>
|
||
</taxonomicName>
|
||
(female) and
|
||
<taxonomicName id="4C034D462853FFC4F359FE3EFB7DB1EE" box="[1086,1246,441,464]" class="Insecta" family="Papilionidae" genus="Hypermnestra" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="genus">
|
||
<emphasis id="B977EAD72853FFC4F359FE3EFB7DB1EE" box="[1086,1246,441,464]" italics="true" pageId="19" pageNumber="20">Hypermnestra</emphasis>
|
||
</taxonomicName>
|
||
(underside), and less clearly, in
|
||
<taxonomicName id="4C034D462853FFC4F621FE5BFE73B1CB" box="[326,464,476,501]" class="Insecta" family="Papilionidae" genus="Allancastria" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="genus">
|
||
<emphasis id="B977EAD72853FFC4F621FE5BFE73B1CB" box="[326,464,476,501]" italics="true" pageId="19" pageNumber="20">Allancastria</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C034D462853FFC4F56AFE5BFDD9B1CB" box="[525,634,476,501]" class="Insecta" family="Papilionidae" genus="Zerynthia" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="genus">
|
||
<emphasis id="B977EAD72853FFC4F56AFE5BFDD9B1CB" box="[525,634,476,501]" italics="true" pageId="19" pageNumber="20">Zerynthia</emphasis>
|
||
</taxonomicName>
|
||
. Some elements of the band, namely a bar or irregular spot over the discocellular veins and a spot on the costa, are probably plesiomorphic within the
|
||
<taxonomicName id="4C034D462853FFC4F32DFE78FB79B226" box="[1098,1242,511,536]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="subFamily" subFamily="Parnassiinae">Parnassiinae</taxonomicName>
|
||
. In that case, the well-defined band of
|
||
<taxonomicName id="4C034D462853FFC4F6E0FDA2FE48B202" box="[391,491,549,572]" class="Insecta" family="Papilionidae" genus="Doritites" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="genus">
|
||
<emphasis id="B977EAD72853FFC4F6E0FDA2FE48B202" box="[391,491,549,572]" italics="true" pageId="19" pageNumber="20">Doritites</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C034D462853FFC4F546FDA3FD3EB203" box="[545,669,548,573]" class="Insecta" family="Papilionidae" genus="Luehdorfia" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="genus">
|
||
<emphasis id="B977EAD72853FFC4F546FDA3FD3EB203" box="[545,669,548,573]" italics="true" pageId="19" pageNumber="20">Luehdorfia</emphasis>
|
||
</taxonomicName>
|
||
could well be synapomorphic.
|
||
</paragraph>
|
||
<paragraph id="8BBC36C52853FFC4F7A0FDC0FC84B3BE" blockId="19.[151,1436,151,1509]" pageId="19" pageNumber="20">
|
||
<bibRefCitation id="EF924B342853FFC4F7A0FDC0FEFDB25E" author="Rebel" box="[199,350,583,608]" pageId="19" pageNumber="20" refString="Rebel, H. (1898) Fossile Lepidopteren aus der Miocan-Formation von Gabbro. Sitzungsberichte der Akademie der Wissenschaften Wien, 107, 731 - 745." type="journal article" year="1898">Rebel (1898)</bibRefCitation>
|
||
thought the fossil to be closely related to the genus
|
||
<taxonomicName id="4C034D462853FFC4F4B3FDCEFB8BB25E" box="[980,1064,585,608]" class="Aves" family="Paridae" genus="Dorites" higherTaxonomySource="GBIF" kingdom="Animalia" order="Passeriformes" pageId="19" pageNumber="20" phylum="Chordata" rank="genus">
|
||
<emphasis id="B977EAD72853FFC4F4B3FDCEFB8BB25E" box="[980,1064,585,608]" italics="true" pageId="19" pageNumber="20">Dorites</emphasis>
|
||
</taxonomicName>
|
||
(now
|
||
<taxonomicName id="4C034D462853FFC4F31FFDC0FB6AB25E" box="[1144,1225,583,608]" class="Insecta" family="Papilionidae" genus="Archon" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="genus">
|
||
<emphasis id="B977EAD72853FFC4F31FFDC0FB6AB25E" box="[1144,1225,583,608]" italics="true" pageId="19" pageNumber="20">Archon</emphasis>
|
||
</taxonomicName>
|
||
), hence the name
|
||
<taxonomicName id="4C034D462853FFC4F7F0FDEAFF58B2BA" box="[151,251,621,644]" class="Insecta" family="Papilionidae" genus="Doritites" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="genus">
|
||
<emphasis id="B977EAD72853FFC4F7F0FDEAFF58B2BA" box="[151,251,621,644]" italics="true" pageId="19" pageNumber="20">Doritites</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
<bibRefCitation id="EF924B342853FFC4F66FFDEAFE11B2BB" author="Munroe" box="[264,434,620,645]" pageId="19" pageNumber="20" refString="Munroe, E. (1961) The classification of the Papilionidae. Canadian Entomologist, 17 (Supplement), 1 - 51." type="journal article" year="1961">Munroe (1961)</bibRefCitation>
|
||
assigned the fossil to
|
||
<taxonomicName id="4C034D462853FFC4F5CEFDEBFC86B2BB" box="[681,805,620,645]" class="Insecta" family="Papilionidae" genus="Luehdorfia" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="genus">
|
||
<emphasis id="B977EAD72853FFC4F5CEFDEBFC86B2BB" box="[681,805,620,645]" italics="true" pageId="19" pageNumber="20">Luehdorfia</emphasis>
|
||
</taxonomicName>
|
||
, as did
|
||
<bibRefCitation id="EF924B342853FFC4F41BFDEBFBA6B2BA" author="Bryk" box="[892,1029,620,645]" pageId="19" pageNumber="20" refString="Bryk, F. (1913) Uber eine neue Einteilung der Papilionidae unter Berucksichtigung des Flugelgeaders. Archiv fur Naturgeschichte, 79 (A 2), 116 - 121." type="journal article" year="1913">Bryk (1913)</bibRefCitation>
|
||
half a century earlier. In view of the discussion above, a relationship of the fossil with the tribe
|
||
<taxonomicName id="4C034D462853FFC4F434FD08FC43B296" box="[851,992,655,680]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="tribe" tribe="Luehdorfiini">Luehdorfiini</taxonomicName>
|
||
(consisting of the genera
|
||
<taxonomicName id="4C034D462853FFC4F270FD08FACBB296" box="[1303,1384,655,680]" class="Insecta" family="Papilionidae" genus="Archon" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="genus">
|
||
<emphasis id="B977EAD72853FFC4F270FD08FACBB296" box="[1303,1384,655,680]" italics="true" pageId="19" pageNumber="20">Archon</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C034D462853FFC4F7F0FD33FEB0B2F3" box="[151,275,692,717]" class="Insecta" family="Papilionidae" genus="Luehdorfia" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="genus">
|
||
<emphasis id="B977EAD72853FFC4F7F0FD33FEB0B2F3" box="[151,275,692,717]" italics="true" pageId="19" pageNumber="20">Luehdorfia</emphasis>
|
||
</taxonomicName>
|
||
) seems supported by the evidence.
|
||
<bibRefCitation id="EF924B342853FFC4F584FD33FC00B2F2" author="Hancock" box="[739,931,692,717]" pageId="19" pageNumber="20" refString="Hancock, D. L. (1983) Classification of the Papilionidae (Lepidoptera): a phylogenetic approach. Smithersia, 2, 1 - 48." type="journal article" year="1983">Hancock (1983)</bibRefCitation>
|
||
considered
|
||
<taxonomicName id="4C034D462853FFC4F327FD32FB07B2F2" box="[1088,1188,693,716]" class="Insecta" family="Papilionidae" genus="Doritites" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="genus">
|
||
<emphasis id="B977EAD72853FFC4F327FD32FB07B2F2" box="[1088,1188,693,716]" italics="true" pageId="19" pageNumber="20">Doritites</emphasis>
|
||
</taxonomicName>
|
||
an offshoot of the
|
||
<taxonomicName id="4C034D462853FFC4F7F0FD50FE81B2CE" box="[151,290,727,752]" class="Insecta" family="Papilionidae" genus="Allancastria" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="genus">
|
||
<emphasis id="B977EAD72853FFC4F7F0FD50FE81B2CE" box="[151,290,727,752]" italics="true" pageId="19" pageNumber="20">Allancastria</emphasis>
|
||
</taxonomicName>
|
||
/
|
||
<taxonomicName id="4C034D462853FFC4F64EFD50FE35B2CE" box="[297,406,727,752]" class="Insecta" family="Papilionidae" genus="Zerynthia" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="genus">
|
||
<emphasis id="B977EAD72853FFC4F64EFD50FE35B2CE" box="[297,406,727,752]" italics="true" pageId="19" pageNumber="20">Zerynthia</emphasis>
|
||
</taxonomicName>
|
||
ancestral line (i.e.
|
||
<taxonomicName id="4C034D462853FFC4F509FD50FD53B2CE" box="[622,752,727,752]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="tribe" tribe="Zerynthiini">Zerynthiini</taxonomicName>
|
||
), but did not give supporting characters.
|
||
<bibRefCitation id="EF924B342853FFC4F3D9FD50FA3FB2CE" box="[1214,1436,727,752]" pageId="19" pageNumber="20" refString="Nazari, V., Zakharov, E. V. & Sperling, F. A. H. (2007) Phylogeny, historical biogeography, and taxonomic ranking of Parnassiinae (Lepidoptera: Papilionidae) based on morphology and seven genes. Molecular Phylogenetics and Evolution, 42, 131 - 156." type="journal article">
|
||
Nazari
|
||
<emphasis id="B977EAD72853FFC4F276FD5EFAE1B2CE" box="[1297,1346,727,752]" italics="true" pageId="19" pageNumber="20">et al</emphasis>
|
||
. (2007)
|
||
</bibRefCitation>
|
||
included
|
||
<taxonomicName id="4C034D462853FFC4F664FD7AFEC4B32A" box="[259,359,765,788]" class="Insecta" family="Papilionidae" genus="Doritites" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="genus">
|
||
<emphasis id="B977EAD72853FFC4F664FD7AFEC4B32A" box="[259,359,765,788]" italics="true" pageId="19" pageNumber="20">Doritites</emphasis>
|
||
</taxonomicName>
|
||
in their analysis, where it ended as sister taxon of
|
||
<taxonomicName id="4C034D462853FFC4F4ACFD7BFBBEB32B" box="[971,1053,764,789]" class="Insecta" family="Papilionidae" genus="Archon" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="genus">
|
||
<emphasis id="B977EAD72853FFC4F4ACFD7BFBBEB32B" box="[971,1053,764,789]" italics="true" pageId="19" pageNumber="20">Archon</emphasis>
|
||
</taxonomicName>
|
||
. At this position they used it as calibration point. As discussed under
|
||
<taxonomicName id="4C034D462853FFC4F523FC98FC94B306" box="[580,823,799,824]" class="Insecta" family="Papilionidae" genus="Praepapilio" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="species" species="colorado">
|
||
<emphasis id="B977EAD72853FFC4F523FC98FC94B306" box="[580,823,799,824]" italics="true" pageId="19" pageNumber="20">Praepapilio colorado</emphasis>
|
||
</taxonomicName>
|
||
, such an inclusion in the analysis must be rejected. If used for calibration purposes, the fossil should be placed at the ancestral line of the
|
||
<taxonomicName id="4C034D462853FFC4F3F9FCC3FA8DB363" box="[1182,1326,836,861]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="tribe" tribe="Luehdorfiini">Luehdorfiini</taxonomicName>
|
||
, a closer relationship with
|
||
<taxonomicName id="4C034D462853FFC4F63CFCE0FE0FB3BE" box="[347,428,871,896]" class="Insecta" family="Papilionidae" genus="Archon" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="genus">
|
||
<emphasis id="B977EAD72853FFC4F63CFCE0FE0FB3BE" box="[347,428,871,896]" italics="true" pageId="19" pageNumber="20">Archon</emphasis>
|
||
</taxonomicName>
|
||
not being supported by evidence.
|
||
</paragraph>
|
||
<paragraph id="8BBC36C52853FFC4F7A0FC0BFA29B40B" blockId="19.[151,1436,151,1509]" pageId="19" pageNumber="20">
|
||
The colours and hindwing tail of
|
||
<taxonomicName id="4C034D462853FFC4F52BFC0AFD13B39A" box="[588,688,909,932]" class="Insecta" family="Papilionidae" genus="Doritites" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="genus">
|
||
<emphasis id="B977EAD72853FFC4F52BFC0AFD13B39A" box="[588,688,909,932]" italics="true" pageId="19" pageNumber="20">Doritites</emphasis>
|
||
</taxonomicName>
|
||
depicted by
|
||
<bibRefCitation id="EF924B342853FFC4F42FFC0BFBDBB39B" author="Bryk" box="[840,1144,908,933]" pageId="19" pageNumber="20" refString="Bryk, F. (1914) Parnassius apollo und sein Formenkreis. Verlagsbuchhandlung R. Stricker, Berlin, 148 pp." type="book" year="1914">Bryk (1914: Pl. 6 Fig. 46)</bibRefCitation>
|
||
were invented by him to make the relationship to
|
||
<taxonomicName id="4C034D462853FFC4F6AFFC28FDE6B3F6" box="[456,581,943,968]" class="Insecta" family="Papilionidae" genus="Luehdorfia" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="genus">
|
||
<emphasis id="B977EAD72853FFC4F6AFFC28FDE6B3F6" box="[456,581,943,968]" italics="true" pageId="19" pageNumber="20">Luehdorfia</emphasis>
|
||
</taxonomicName>
|
||
seems more plausible. Later (
|
||
<bibRefCitation id="EF924B342853FFC4F4DFFC28FBDBB3F6" author="Bryk" box="[952,1144,943,968]" pageId="19" pageNumber="20" refString="Bryk, F. (1934) Baroniidae, Teinopalpidae, Parnassiidae pars 1. In: Das Tierreich 64. De Gruyter, Berlin, Leipzig, xxiii + 790 pp." type="book" year="1934">Bryk 1934: 112</bibRefCitation>
|
||
) he wrote: “[...] ob der abweichende Praecostalsporn des Hinterflügels, der einfach distal gekrümmt und nicht 2ästig ist, tatsächlich so gebaut war oder unrichtig rekonstruiert wurde, sei dahingestellt.” ("[…] whether the deviating precostal thorn of the hindwing really was simply curved and not branched, or was reconstructed incorrectly, remains undecided.").
|
||
</paragraph>
|
||
<paragraph id="8BBC36C52853FFC4F7A0FBB8FBE7B533" blockId="19.[151,1436,151,1509]" pageId="19" pageNumber="20">
|
||
References with regard to this fossil up to 1934 can be found in
|
||
<bibRefCitation id="EF924B342853FFC4F4E8FBB8FBB8B466" author="Bryk" box="[911,1051,1087,1112]" pageId="19" pageNumber="20" refString="Bryk, F. (1934) Baroniidae, Teinopalpidae, Parnassiidae pars 1. In: Das Tierreich 64. De Gruyter, Berlin, Leipzig, xxiii + 790 pp." type="book" year="1934">Bryk (1934)</bibRefCitation>
|
||
. A single paper may be mentioned here, that of
|
||
<bibRefCitation id="EF924B342853FFC4F64EFBE3FE60B442" author="Turati" box="[297,451,1124,1149]" pageId="19" pageNumber="20" refString="Turati, E. (1918) Variabilita del Parnassius apollo pumilus Stich. a ricerche sull' origine dei Parnassius. Atti Societa Italiana di Scienza naturali, 57, 29 - 89." type="journal article" year="1918">Turati (1918)</bibRefCitation>
|
||
. This author considered
|
||
<taxonomicName id="4C034D462853FFC4F5BCFBE2FC12B443" box="[731,945,1124,1149]" class="Insecta" family="Papilionidae" genus="Doritites" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="species" species="bosnaskii">
|
||
<emphasis id="B977EAD72853FFC4F5BCFBE2FC12B443" box="[731,945,1124,1149]" italics="true" pageId="19" pageNumber="20">Doritites bosnaskii</emphasis>
|
||
</taxonomicName>
|
||
the ancestor of two subspecies of different
|
||
<taxonomicName id="4C034D462853FFC4F7F0FB0EFEB0B49E" box="[151,275,1161,1184]" class="Insecta" family="Papilionidae" genus="Parnassius" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="genus">
|
||
<emphasis id="B977EAD72853FFC4F7F0FB0EFEB0B49E" box="[151,275,1161,1184]" italics="true" pageId="19" pageNumber="20">Parnassius</emphasis>
|
||
</taxonomicName>
|
||
species:
|
||
<emphasis id="B977EAD72853FFC4F61BFB00FD86B49E" box="[380,549,1159,1184]" italics="true" pageId="19" pageNumber="20">
|
||
<taxonomicName id="4C034D462853FFC4F61BFB00FE67B49E" box="[380,452,1159,1184]" class="Insecta" family="Papilionidae" genus="Parnassius" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="species" species="apollo">apollo</taxonomicName>
|
||
pumilus
|
||
</emphasis>
|
||
Stichel and
|
||
<emphasis id="B977EAD72853FFC4F5D4FB0EFC17B49E" box="[691,948,1159,1184]" italics="true" pageId="19" pageNumber="20">mnemosyne calabricus</emphasis>
|
||
Turati. More recent opinions can be found in
|
||
<bibRefCitation id="EF924B342853FFC4F7D2FB2BFEEEB4FA" author="Verity" box="[181,333,1196,1221]" pageId="19" pageNumber="20" refString="Verity, R. (1947) Le farfalle diurne d'Italia. Fol. 3. Marzocco, Firenze, xvi + 318 pp." type="book" year="1947">Verity (1947)</bibRefCitation>
|
||
who, without much comment, assigns the fossil to “Zerynthiinae-Lühdorfiidi”, and in
|
||
<bibRefCitation id="EF924B342853FFC4F24BFB2BFF42B4D6" author="Demoulin" pageId="19" pageNumber="20" refString="Demoulin, G. (1975) A propos de deux Papilionidae mediterraneens fossiles. Bulletin Cercle des Lepidopteristes de Belgique, 4, 96 - 101." type="journal article" year="1975">Demoulin (1975)</bibRefCitation>
|
||
who followed Bryk's (1914) suggestion. With or without valid arguments, recent authors are thus unanimous in assigning the fossil to the Palaearctic tribes
|
||
<taxonomicName id="4C034D462853FFC4F447FB73FC03B533" box="[800,928,1268,1293]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="tribe" tribe="Zerynthiini">Zerynthiini</taxonomicName>
|
||
+
|
||
<taxonomicName id="4C034D462853FFC4F4C9FB73FB9CB533" box="[942,1087,1268,1293]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="tribe" tribe="Luehdorfiini">Luehdorfiini</taxonomicName>
|
||
.
|
||
</paragraph>
|
||
<paragraph id="8BBC36C52853FFC4F7A0FA9FFA96B5A3" blockId="19.[151,1436,151,1509]" pageId="19" pageNumber="20">
|
||
Larsen’s (1974: 34) remark that “
|
||
<taxonomicName id="4C034D462853FFC4F52CFA9EFD64B50E" box="[587,711,1305,1328]" class="Insecta" family="Papilionidae" genus="Parnassius" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="genus">
|
||
<emphasis id="B977EAD72853FFC4F52CFA9EFD64B50E" box="[587,711,1305,1328]" italics="true" pageId="19" pageNumber="20">Parnassius</emphasis>
|
||
</taxonomicName>
|
||
-like butterflies, close to
|
||
<taxonomicName id="4C034D462853FFC4F48EFA90FB0FB50E" box="[1001,1196,1303,1328]" class="Insecta" family="Papilionidae" genus="Archon" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="species" species="apollinus">
|
||
<emphasis id="B977EAD72853FFC4F48EFA90FB0FB50E" box="[1001,1196,1303,1328]" italics="true" pageId="19" pageNumber="20">Archon apollinus</emphasis>
|
||
</taxonomicName>
|
||
, have been found in Baltic Amber” and uncritically quoted by
|
||
<bibRefCitation id="EF924B342853FFC4F51CFABBFC82B56B" author="Shields" box="[635,801,1340,1365]" pageId="19" pageNumber="20" refString="Shields, O. (1976) Fossil butterflies and the evolution of Lepidoptera. Journal of Research on the Lepidoptera, 15, 132 - 143." type="journal article" year="1976">Shields (1976)</bibRefCitation>
|
||
must be based on a misunderstanding.
|
||
<bibRefCitation id="EF924B342853FFC4F38EFABAFA34B56A" author="Larsson" box="[1257,1431,1340,1364]" pageId="19" pageNumber="20" refString="Larsson, S. G. (1978) Baltic Amber - a Palaeobiological Study. Scandinavian Science Press, Klampenborg, 192 pp." type="book" year="1978">Larsson (1978)</bibRefCitation>
|
||
, while dealing with all insects in Baltic Amber, does not mention any butterfly. Larsen’s remark must refer to
|
||
<taxonomicName id="4C034D462853FFC4F2E6FAE6FEAAB5A3" class="Insecta" family="Papilionidae" genus="Doritites" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="species" species="bosniaskii">
|
||
<emphasis id="B977EAD72853FFC4F2E6FAE6FEAAB5A3" italics="true" pageId="19" pageNumber="20">D. bosniaskii</emphasis>
|
||
</taxonomicName>
|
||
or to
|
||
<taxonomicName id="4C034D462853FFC4F62DFA03FDBAB5A2" box="[330,537,1412,1437]" class="Insecta" family="Papilionidae" genus="Thaites" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="species" species="ruminiana">
|
||
<emphasis id="B977EAD72853FFC4F62DFA03FDBAB5A2" box="[330,537,1412,1437]" italics="true" pageId="19" pageNumber="20">Thaites ruminiana</emphasis>
|
||
</taxonomicName>
|
||
, the only
|
||
<taxonomicName id="4C034D462853FFC4F5EFFA02FCA7B5A2" box="[648,772,1413,1436]" class="Insecta" family="Papilionidae" genus="Parnassius" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="genus">
|
||
<emphasis id="B977EAD72853FFC4F5EFFA02FCA7B5A2" box="[648,772,1413,1436]" italics="true" pageId="19" pageNumber="20">Parnassius</emphasis>
|
||
</taxonomicName>
|
||
-like fossil butterflies known, but not from amber.
|
||
</paragraph>
|
||
<paragraph id="8BBC36C52853FFC4F7A0FA20FDF1B5DA" blockId="19.[151,1436,151,1509]" pageId="19" pageNumber="20">
|
||
With two other papilionid fossils, this fossil was used for calibration by
|
||
<bibRefCitation id="EF924B342853FFC4F359FA20FAC5B5FE" box="[1086,1382,1447,1472]" pageId="19" pageNumber="20" refString="Condamine, F. L., Sperling, F. A. H., Wahlberg, N., Rasplus, J. - Y. & Kergoat, G. (2012) What causes latitudunal gradients in species diversity? Evolutionary processes and ecological constraints on swallowtail biodiversity. Ecology Letters, 15, 267 - 277." type="journal article">
|
||
Condamine
|
||
<emphasis id="B977EAD72853FFC4F3ABFA2EFAAFB5FE" box="[1228,1292,1447,1472]" italics="true" pageId="19" pageNumber="20">et al.</emphasis>
|
||
(2012)
|
||
</bibRefCitation>
|
||
, see discussion under
|
||
<taxonomicName id="4C034D462853FFC4F63EFA4BFDE8B5DB" box="[345,587,1484,1509]" class="Insecta" family="Papilionidae" genus="Praepapilio" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="species" species="colorado">
|
||
<emphasis id="B977EAD72853FFC4F63EFA4BFDE8B5DB" box="[345,587,1484,1509]" italics="true" pageId="19" pageNumber="20">Praepapilio colorado</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |