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<mods:title id="7144EAF45A5331AF3F2159775411E1BF">Fossil butterflies, calibration points and the molecular clock (Lepidoptera: Papilionoidea)</mods:title>
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<treatment id="03AA87D32863FFF3F7F0F9DCFF4CB1F1" ID-DOI="http://doi.org/10.5281/zenodo.6047038" ID-GBIF-Taxon="130844905" ID-Zenodo-Dep="6047038" LSID="urn:lsid:plazi:treatment:03AA87D32863FFF3F7F0F9DCFF4CB1F1" httpUri="http://treatment.plazi.org/id/03AA87D32863FFF3F7F0F9DCFF4CB1F1" lastPageId="36" lastPageNumber="37" pageId="35" pageNumber="36">
<subSubSection id="C319654E2863FFF4F7F0F9DCFE0DB684" pageId="35" pageNumber="36" type="nomenclature">
<paragraph id="8BBC36C52863FFF4F7F0F9DCFD2BB64B" blockId="35.[151,648,1627,1653]" box="[151,648,1627,1653]" pageId="35" pageNumber="36">
<heading id="D0F481A92863FFF4F7F0F9DCFD2BB64B" bold="true" box="[151,648,1627,1653]" fontSize="11" level="1" pageId="35" pageNumber="36" reason="1">
<emphasis id="B977EAD72863FFF4F7F0F9DCFD2BB64B" bold="true" box="[151,648,1627,1653]" pageId="35" pageNumber="36">
<emphasis id="B977EAD72863FFF4F7F0F9DCFF5EB64B" bold="true" box="[151,253,1627,1653]" italics="true" pageId="35" pageNumber="36">
<taxonomicName id="4C034D462863FFF4F7F0F9DCFF5BB64B" box="[151,248,1627,1653]" class="Insecta" family="Nymphalidae" genus="Satyrites" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="35" pageNumber="36" phylum="Arthropoda" rank="species" species="reynesii">reynesii</taxonomicName>
.
</emphasis>
<taxonomicName id="4C034D462863FFF4F663F9DCFD2BB64B" authority="Scudder, 1872" authorityName="Scudder" authorityYear="1872" box="[260,648,1627,1653]" class="Insecta" family="Nymphalidae" genus="Satyrites" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="35" pageNumber="36" phylum="Arthropoda" rank="species" species="reynesii">
<emphasis id="B977EAD72863FFF4F663F9DCFE72B64B" bold="true" box="[260,465,1627,1653]" italics="true" pageId="35" pageNumber="36">Satyrites reynesii</emphasis>
<bibRefCitation id="EF924B342863FFF4F6BEF9DCFD2BB64B" author="Scudder" box="[473,648,1627,1653]" pageId="35" pageNumber="36" refString="Scudder, S. H. (1872) Description d'un nouveau papillon fossile (Satyrites Reynesii) trouve a Aix en Provence. Revue et Magasin de Zoologie, 23, (2), 66 - 72. cudder, S. H. (1875) Fossil butterflies. Memoirs American Association Advancement Science, 1, 1 - 99." type="journal article" year="1872">Scudder, 1872</bibRefCitation>
</taxonomicName>
</emphasis>
</heading>
</paragraph>
<paragraph id="8BBC36C52863FFF4F7F0F926FE0DB684" blockId="35.[151,1437,1697,2010]" box="[151,430,1697,1722]" pageId="35" pageNumber="36">
<taxonomicName id="4C034D462863FFF4F7F0F926FE90B684" box="[151,307,1697,1722]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="35" pageNumber="36" phylum="Arthropoda" rank="family">Nymphalidae</taxonomicName>
:
<taxonomicName id="4C034D462863FFF4F65AF926FE08B684" box="[317,427,1697,1722]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="35" pageNumber="36" phylum="Arthropoda" rank="subFamily" subFamily="Satyrinae">Satyrinae</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection id="C319654E2863FFF3F7A0F941FF4CB1F1" lastPageId="36" lastPageNumber="37" pageId="35" pageNumber="36" type="description">
<paragraph id="8BBC36C52863FFF4F7A0F941FB8EB6E3" blockId="35.[151,1437,1697,2010]" box="[199,1069,1733,1758]" pageId="35" pageNumber="36">France, Bouches-du-Rhône, Aix-en-Provence; late Oligoceneearly Miocene.</paragraph>
<paragraph id="8BBC36C52863FFF4F7A0F96EFD88B73C" blockId="35.[151,1437,1697,2010]" box="[199,555,1769,1794]" pageId="35" pageNumber="36">Depository: MVMF (holotype).</paragraph>
<paragraph id="8BBC36C52863FFF4F7A0F88AFBEAB718" blockId="35.[151,1437,1697,2010]" box="[199,1097,1805,1830]" pageId="35" pageNumber="36">
Published figures:
<bibRefCitation id="EF924B342863FFF4F6FEF889FD47B718" author="Leestmans" box="[409,740,1805,1830]" pageId="35" pageNumber="36" refString="Leestmans, R. (1983) Les Lepidopteres fossiles trouves en France (Insecta Lepidoptera). Linneana belgica, 9, 64 - 89." type="journal article" year="1983">Leestmans (1983: Figs 910)</bibRefCitation>
; Scudder (1875: Pl. I
<figureCitation id="13382A402863FFF4F4BFF88AFBBCB718" box="[984,1055,1805,1830]" 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="35" pageNumber="36">Figs 2</figureCitation>
,
<figureCitation id="13382A402863FFF4F34CF88AFB98B718" box="[1067,1083,1805,1830]" captionStart="FIGURE 5" captionStartId="16.[151,250,1784,1806]" captionTargetBox="[238,1332,193,1743]" captionTargetId="figure@16.[238,1332,193,1754]" captionTargetPageId="16" captionText="FIGURE 5. Ƒanessa amerindica Miller &amp; Brown, 1989, holotype, collections of Invertebrate Paleontology, University of Florida. Photo by Roger Portell. Fig. 6. Pseudoneorina coulleti Nel &amp; 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 &amp; 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="https://zenodo.org/record/583187/files/figure.png" pageId="35" pageNumber="36">5</figureCitation>
).
</paragraph>
<paragraph id="8BBC36C52863FFF3F7A0F8B6FB42B17E" blockId="35.[151,1437,1697,2010]" lastBlockId="36.[151,1437,151,464]" lastPageId="36" lastPageNumber="37" pageId="35" pageNumber="36">
The original description (
<bibRefCitation id="EF924B342863FFF4F686F8B6FDDCB774" author="Scudder" box="[481,639,1841,1866]" pageId="35" pageNumber="36" refString="Scudder, S. H. (1872) Description d'un nouveau papillon fossile (Satyrites Reynesii) trouve a Aix en Provence. Revue et Magasin de Zoologie, 23, (2), 66 - 72. cudder, S. H. (1875) Fossil butterflies. Memoirs American Association Advancement Science, 1, 1 - 99." type="journal article" year="1872">Scudder 1872</bibRefCitation>
) is repeated by Scudder (1875), who erected a new genus for the fossil:
<taxonomicName id="4C034D462863FFF4F7F0F8D2FF53B750" box="[151,240,1877,1902]" class="Insecta" family="Nymphalidae" genus="Lethites" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="35" pageNumber="36" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72863FFF4F7F0F8D2FF53B750" box="[151,240,1877,1902]" italics="true" pageId="35" pageNumber="36">Lethites</emphasis>
</taxonomicName>
.
<bibRefCitation id="EF924B342863FFF4F661F8D2FE10B750" author="Kozlov" box="[262,435,1877,1902]" pageId="35" pageNumber="36" refString="Kozlov, M. V. (1988) Paleontology of lepidopterans and problems of the phylogeny of the order Papilionida. In: Ponomarenko, A. G. (Ed.), The Mesozoic-Cenozoic Crisis in the Evolution of Insects. Academy of Sciences, Moscow, pp. 16 - 69. [in Russian]" type="book chapter" year="1988">Kozlov (1988)</bibRefCitation>
accepted Scudders genus switch. The poorly preserved specimen bears only the forewings, which yield some usable characters. Subcosta basally strongly swollen; radial formula 1, 2, 3+(4+5), cell closed. A swollen subcosta is typical, but not universal, for
<taxonomicName id="4C034D462863FFF4F404F81AFB23B788" box="[867,1152,1949,1974]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="35" pageNumber="36" phylum="Arthropoda" rank="subFamily" subFamily="Satyrinae">Satyrinae (Nymphalidae)</taxonomicName>
, but this character occurs in some genera of the
<taxonomicName id="4C034D462863FFF4F6F0F846FDAFB7E4" box="[407,524,1985,2010]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="35" pageNumber="36" phylum="Arthropoda" rank="subFamily" subFamily="Biblidinae">Biblidinae</taxonomicName>
as well (
<taxonomicName id="4C034D462863FFF4F511F846FD14B7E4" box="[630,695,1985,2010]" class="Insecta" family="Nymphalidae" genus="Biblis" kingdom="Animalia" order="Lepidoptera" pageId="35" pageNumber="36" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72863FFF4F511F846FD14B7E4" box="[630,695,1985,2010]" italics="true" pageId="35" pageNumber="36">Biblis</emphasis>
</taxonomicName>
,
<taxonomicName id="4C034D462863FFF4F5A0F846FCBBB7E4" box="[711,792,1985,2010]" class="Insecta" family="Nymphalidae" genus="Ergolis" higherTaxonomySource="CoL" kingdom="Animalia" order="Lepidoptera" pageId="35" pageNumber="36" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72863FFF4F5A0F846FCBBB7E4" box="[711,792,1985,2010]" italics="true" pageId="35" pageNumber="36">Ergolis</emphasis>
</taxonomicName>
,
<taxonomicName id="4C034D462863FFF4F44FF846FC2BB7E4" box="[808,904,1985,2010]" class="Insecta" family="Nymphalidae" genus="Eurytela" kingdom="Animalia" order="Lepidoptera" pageId="35" pageNumber="36" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72863FFF4F44FF846FC2BB7E4" box="[808,904,1985,2010]" italics="true" pageId="35" pageNumber="36">Eurytela</emphasis>
</taxonomicName>
,
<taxonomicName id="4C034D462863FFF4F4FFF845FC56B7E7" box="[920,1013,1986,2009]" class="Insecta" family="Nymphalidae" genus="Hypanis" kingdom="Animalia" order="Lepidoptera" pageId="35" pageNumber="36" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72863FFF4F4FFF845FC56B7E7" box="[920,1013,1986,2009]" italics="true" pageId="35" pageNumber="36">Hypanis</emphasis>
</taxonomicName>
,
<taxonomicName id="4C034D462863FFF4F362F846FB92B7E4" box="[1029,1073,1985,2010]" class="Insecta" family="Nymphalidae" genus="Vila" kingdom="Animalia" order="Lepidoptera" pageId="35" pageNumber="36" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72863FFF4F362F846FB92B7E4" box="[1029,1073,1985,2010]" italics="true" pageId="35" pageNumber="36">Vila</emphasis>
</taxonomicName>
). In
<taxonomicName id="4C034D462863FFF4F30CF846FAA1B7E4" box="[1131,1282,1985,2010]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="35" pageNumber="36" phylum="Arthropoda" rank="subFamily" subFamily="Nymphalinae">Nymphalinae</taxonomicName>
with swollen Sc, the cell is usually open or at least, the ldc is weak. In the fossil ldc seems to be fairly strong. Thus, it is plausible to consider the fossil a member of the
<taxonomicName id="4C034D462864FFF3F52DFF3BFD14B0EB" box="[586,695,188,213]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="36" pageNumber="37" phylum="Arthropoda" rank="subFamily" subFamily="Satyrinae">Satyrinae</taxonomicName>
. According to Scudder it is similar to the recent genera
<taxonomicName id="4C034D462864FFF3F24AFF3BFACFB0EB" box="[1325,1388,188,213]" class="Insecta" family="Nymphalidae" genus="Lethe" kingdom="Animalia" order="Lepidoptera" pageId="36" pageNumber="37" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72864FFF3F24AFF3BFACFB0EB" box="[1325,1388,188,213]" italics="true" pageId="36" pageNumber="37">Lethe</emphasis>
</taxonomicName>
and
<taxonomicName id="4C034D462864FFF3F7F0FF67FEABB0C9" box="[151,264,224,247]" class="Insecta" family="Nymphalidae" genus="Cercyonis" kingdom="Animalia" order="Lepidoptera" pageId="36" pageNumber="37" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72864FFF3F7F0FF67FEABB0C9" box="[151,264,224,247]" italics="true" pageId="36" pageNumber="37">Cercyonis</emphasis>
</taxonomicName>
, but it is not clear why it should be more similar to these genera than to any other satyrine genus. The similarity is not an indication of relationship, since
<taxonomicName id="4C034D462864FFF3F5BEFE83FCBBB123" box="[729,792,260,285]" class="Insecta" family="Nymphalidae" genus="Lethe" kingdom="Animalia" order="Lepidoptera" pageId="36" pageNumber="37" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72864FFF3F5BEFE83FCBBB123" box="[729,792,260,285]" italics="true" pageId="36" pageNumber="37">Lethe</emphasis>
</taxonomicName>
and
<taxonomicName id="4C034D462864FFF3F428FE82FC1CB122" box="[847,959,261,284]" class="Insecta" family="Nymphalidae" genus="Cercyonis" kingdom="Animalia" order="Lepidoptera" pageId="36" pageNumber="37" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72864FFF3F428FE82FC1CB122" box="[847,959,261,284]" italics="true" pageId="36" pageNumber="37">Cercyonis</emphasis>
</taxonomicName>
are only remotely related (
<bibRefCitation id="EF924B342864FFF3F388FE83FF77B17E" author="Wahlberg" pageId="36" pageNumber="37" refString="Wahlberg, N., Leneveu, J., Kodandaramaiah, U., Pena, C., Nylin, S., Freitas, A. V. L. &amp; Brower, A. V. Z. (2009) Nymphalid butterflies diversify following near demise at the Cretaceous / Tertiary boundary. Proceedings Royal Society B, 276, 4295 - 4302." type="journal article" year="2009">
Wahlberg
<emphasis id="B977EAD72864FFF3F203FE82FA36B123" box="[1380,1429,260,285]" italics="true" pageId="36" pageNumber="37">et al</emphasis>
. 2009
</bibRefCitation>
). Moreover, in
<taxonomicName id="4C034D462864FFF3F619FEA0FE1EB17E" box="[382,445,295,320]" class="Insecta" family="Nymphalidae" genus="Lethe" kingdom="Animalia" order="Lepidoptera" pageId="36" pageNumber="37" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72864FFF3F619FEA0FE1EB17E" box="[382,445,295,320]" italics="true" pageId="36" pageNumber="37">Lethe</emphasis>
</taxonomicName>
there are no basally swollen veins in the forewing (
<bibRefCitation id="EF924B342864FFF3F499FEA0FB77B17E" author="Ackery" box="[1022,1236,295,320]" pageId="36" pageNumber="37" refString="Ackery, P. R., de Jong, R. &amp; 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">
Ackery
<emphasis id="B977EAD72864FFF3F33DFEAEFB29B17E" box="[1114,1162,295,320]" italics="true" pageId="36" pageNumber="37">et al</emphasis>
. 1999
</bibRefCitation>
).
</paragraph>
<paragraph id="8BBC36C52864FFF3F7A0FECAFF4CB1F1" blockId="36.[151,1437,151,464]" pageId="36" pageNumber="37">
<emphasis id="B977EAD72864FFF3F7A0FECAFDA5B15B" box="[199,518,332,357]" italics="true" pageId="36" pageNumber="37">
Neorinopsis,
<taxonomicName id="4C034D462864FFF3F63BFECBFDA5B15B" box="[348,518,332,357]" class="Insecta" family="Nymphalidae" genus="Pseudoneorina" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="36" pageNumber="37" phylum="Arthropoda" rank="genus">Pseudoneorina</taxonomicName>
</emphasis>
,
<taxonomicName id="4C034D462864FFF3F573FECBFD2FB15B" box="[532,652,332,357]" class="Insecta" family="Nymphalidae" genus="Neorinella" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="36" pageNumber="37" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72864FFF3F573FECBFD2FB15B" box="[532,652,332,357]" italics="true" pageId="36" pageNumber="37">Neorinella</emphasis>
</taxonomicName>
and
<taxonomicName id="4C034D462864FFF3F5A4FECAFC86B15A" box="[707,805,333,356]" class="Insecta" family="Nymphalidae" genus="Satyrites" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="36" pageNumber="37" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72864FFF3F5A4FECAFC86B15A" box="[707,805,333,356]" italics="true" pageId="36" pageNumber="37">Satyrites</emphasis>
</taxonomicName>
have been classified, without comment, as fossil genera belonging to the subtribe
<taxonomicName id="4C034D462864FFF3F6A1FEE8FDB8B1B6" box="[454,539,367,392]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="36" pageNumber="37" phylum="Arthropoda" rank="subTribe" subTribe="Lethina">Lethina</taxonomicName>
of the tribe
<taxonomicName id="4C034D462864FFF3F5D1FEE8FCACB1B6" box="[694,783,367,392]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="36" pageNumber="37" phylum="Arthropoda" rank="tribe" tribe="Satyrini">Satyrini</taxonomicName>
by
<bibRefCitation id="EF924B342864FFF3F426FEE8FBCBB1B6" author="Wahlberg" box="[833,1128,367,392]" pageId="36" pageNumber="37" refString="Wahlberg, N. &amp; Pena, C. (2015) Systematics and Evolution of Nymphalidae. Available from: http: // www. nymphalidae. net / (accessed 12 November 2015)" type="journal article" year="2015">Wahlberg &amp; Peña (2015)</bibRefCitation>
, but these choices are not supported by apomorphies. For further details, see the discussion under
<taxonomicName id="4C034D462864FFF3F49EFE13FB71B192" box="[1017,1234,404,429]" class="Insecta" family="Nymphalidae" genus="Neorinella" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="36" pageNumber="37" phylum="Arthropoda" rank="species" species="garciae">
<emphasis id="B977EAD72864FFF3F49EFE13FB71B192" box="[1017,1234,404,429]" italics="true" pageId="36" pageNumber="37">Neorinella garciae</emphasis>
</taxonomicName>
and
<emphasis id="B977EAD72864FFF3F273FE12FF4BB1EE" italics="true" pageId="36" pageNumber="37">
Neorinopsis
<taxonomicName id="4C034D462864FFF3F7F0FE30FF4BB1EE" box="[151,232,439,464]" class="Insecta" family="Nymphalidae" genus="Cyllo" higherTaxonomySource="CoL" kingdom="Animalia" order="Lepidoptera" pageId="36" pageNumber="37" phylum="Arthropoda" rank="species" species="sepulta">sepulta</taxonomicName>
</emphasis>
.
</paragraph>
</subSubSection>
</treatment>
</document>