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<document id="415FA1226B7A5A3FB36A1A1C94B6ED19" ID="10.11646/zootaxa.4270.1.1SLASH11140" ID-CLB-Dataset="32975" ID-DOI="10.5281/zenodo.583183" ID-GBIF-Dataset="6c479acc-8b18-4f0b-a6e5-85bcd6d7b6b7" ID-ISSN="1175-5326" ID-Zenodo-Dep="583183" ID-ZooBank="2D00AFF5-4FE2-4EC1-A328-C8670CFB8D6D" IM.bibliography_approvedBy="felipe" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.taxonomicNames_approvedBy="felipe" checkinTime="1495696370209" checkinUser="plazi" docAuthor="Jong, Rienk De" docDate="2017" docId="03AA87D32854FFC2F7F0FABCFBC3B683" docLanguage="en" docName="zootaxa.4270.1.1SLASH11140.pdf" docOrigin="Zootaxa 4270 (1)" docStyle="DocumentStyle:5EBBA59367AD13919D70D935FA04F6A3.14:Zootaxa.2013-.monograph" docStyleId="5EBBA59367AD13919D70D935FA04F6A3" docStyleName="Zootaxa.2013-.monograph" docStyleVersion="14" docTitle="Praepapilio colorado" docType="treatment" docVersion="9" lastPageNumber="22" masterDocId="FF93FFAB2840FFD7F767FF87FFA3B03E" masterDocTitle="Fossil butterflies, calibration points and the molecular clock (Lepidoptera: Papilionoidea)" masterLastPageNumber="63" masterPageNumber="1" pageNumber="21" updateTime="1698439901205" updateUser="ExternalLinkService">
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<mods:title id="B8765B9F95F8FDBAED9F43416F993E05">Fossil butterflies, calibration points and the molecular clock (Lepidoptera: Papilionoidea)</mods:title>
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<mods:namePart id="9ED246F3AE25A0759C88C7487CE2DDED">Jong, Rienk De</mods:namePart>
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<mods:identifier id="6D88208F00E5979A5B2CC070FC1B9843" type="DOI">10.5281/zenodo.583183</mods:identifier>
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<treatment id="03AA87D32854FFC2F7F0FABCFBC3B683" ID-DOI="http://doi.org/10.5281/zenodo.6046934" ID-GBIF-Taxon="130844890" ID-Zenodo-Dep="6046934" LSID="urn:lsid:plazi:treatment:03AA87D32854FFC2F7F0FABCFBC3B683" httpUri="http://treatment.plazi.org/id/03AA87D32854FFC2F7F0FABCFBC3B683" lastPageId="21" lastPageNumber="22" pageId="20" pageNumber="21">
<subSubSection id="C319654E2854FFC3F7F0FABCFE88B5A4" pageId="20" pageNumber="21" type="nomenclature">
<paragraph id="8BBC36C52854FFC3F7F0FABCFC82B56B" blockId="20.[151,801,1339,1365]" box="[151,801,1339,1365]" pageId="20" pageNumber="21">
<heading id="D0F481A92854FFC3F7F0FABCFC82B56B" bold="true" box="[151,801,1339,1365]" fontSize="11" level="1" pageId="20" pageNumber="21" reason="1">
<emphasis id="B977EAD72854FFC3F7F0FABCFC82B56B" bold="true" box="[151,801,1339,1365]" pageId="20" pageNumber="21">
<emphasis id="B977EAD72854FFC3F7F0FABCFEABB56B" bold="true" box="[151,264,1339,1365]" italics="true" pageId="20" pageNumber="21">
<taxonomicName id="4C034D462854FFC3F7F0FABCFEA7B56B" box="[151,260,1339,1365]" class="Insecta" family="Papilionidae" genus="Praepapilio" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="20" pageNumber="21" phylum="Arthropoda" rank="species" species="colorado">colorado</taxonomicName>
.
</emphasis>
<taxonomicName id="4C034D462854FFC3F668FABCFC82B56B" authority="Durden &amp; Rose, 1978" authorityName="Durden &amp; Rose" authorityYear="1978" box="[271,801,1339,1365]" class="Insecta" family="Papilionidae" genus="Praepapilio" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="20" pageNumber="21" phylum="Arthropoda" rank="species" species="colorado">
<emphasis id="B977EAD72854FFC3F668FABCFDAEB56B" bold="true" box="[271,525,1339,1365]" italics="true" pageId="20" pageNumber="21">Praepapilio colorado</emphasis>
<bibRefCitation id="EF924B342854FFC3F574FABCFC82B56B" author="Durden" box="[531,801,1339,1365]" pageId="20" pageNumber="21" refString="Durden, C. J. &amp; Rose, H. (1978) Butterflies from the Middle Miocene: the earliest occurrence of fossil Papilionoidea (Lepidoptera). Pearce-Sellards Series, 29, 1 - 25." type="journal article" year="1978">Durden &amp; Rose, 1978</bibRefCitation>
</taxonomicName>
</emphasis>
</heading>
</paragraph>
<paragraph id="8BBC36C52854FFC3F7F0FA06FE88B5A4" blockId="20.[151,1437,1409,2010]" box="[151,299,1409,1434]" pageId="20" pageNumber="21">
<taxonomicName id="4C034D462854FFC3F7F0FA06FE84B5A4" box="[151,295,1409,1434]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="20" pageNumber="21" phylum="Arthropoda" rank="family">Papilionidae</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection id="C319654E2854FFC2F7A0FA21FA3FB56A" lastPageId="21" lastPageNumber="22" pageId="20" pageNumber="21" type="description">
<paragraph id="8BBC36C52854FFC3F7A0FA21FBBCB583" blockId="20.[151,1437,1409,2010]" box="[199,1055,1445,1470]" pageId="20" pageNumber="21">
<materialsCitation id="3B6B3C982854FFC3F7A0FA21FBBCB583" ID-GBIF-Occurrence="1503739231" box="[199,1055,1445,1470]" country="United States of America" county="Rio Blanco Co." location="Ray Dome" pageId="20" pageNumber="21" specimenCount="1" stateProvince="Colorado">
<collectingCountry id="F31476552854FFC3F7A0FA21FEA1B580" box="[199,258,1446,1470]" name="United States of America" pageId="20" pageNumber="21">USA</collectingCountry>
,
<taxonomicName id="4C034D462854FFC3F66AFA22FEDBB580" box="[269,376,1445,1470]" class="Insecta" family="Papilionidae" genus="Praepapilio" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="20" pageNumber="63" phylum="Arthropoda" rank="species" species="colorado">
<collectingRegion id="49C7F8272854FFC3F66AFA22FEDBB580" box="[269,376,1445,1470]" country="United States of America" name="Colorado" pageId="20" pageNumber="21">Colorado</collectingRegion>
</taxonomicName>
,
<location id="8EDC601E2854FFC3F6E4FA21FE5DB583" LSID="urn:lsid:plazi:treatment:03AA87D32854FFC2F7F0FABCFBC3B683:8EDC601E2854FFC3F6E4FA21FE5DB583" box="[387,510,1446,1470]" country="United States of America" county="Rio Blanco Co." name="Ray Dome" pageId="20" pageNumber="21" stateProvince="Colorado">Ray Dome</location>
,
<collectingCounty id="62DD4E492854FFC3F56DFA22FD19B580" box="[522,698,1445,1470]" pageId="20" pageNumber="21">Rio Blanco Co.</collectingCounty>
; early Lutetian, middle Eocene.
</materialsCitation>
</paragraph>
<paragraph id="8BBC36C52854FFC3F7A0FA4EFBE0B5DC" blockId="20.[151,1437,1409,2010]" box="[199,1091,1481,1506]" pageId="20" pageNumber="21">
<materialsCitation id="3B6B3C982854FFC3F7A0FA4EFBE0B5DC" ID-GBIF-Occurrence="1503739217" box="[199,1091,1481,1506]" country="United States of America" location="Depository" municipality="Hugh Rose" pageId="20" pageNumber="21" specimenCount="1" stateProvince="New Hampshire" typeStatus="holotype">
<location id="8EDC601E2854FFC3F7A0FA4EFEEBB5DC" LSID="urn:lsid:plazi:treatment:03AA87D32854FFC2F7F0FABCFBC3B683:8EDC601E2854FFC3F7A0FA4EFEEBB5DC" box="[199,328,1481,1506]" country="United States of America" municipality="Hugh Rose" name="Depository" pageId="20" pageNumber="21" stateProvince="New Hampshire">Depository</location>
: private collection
<collectingMunicipality id="6BD8ACBF2854FFC3F57BFA4EFD3FB5DF" box="[540,668,1481,1506]" pageId="20" pageNumber="21">Hugh Rose</collectingMunicipality>
,
<collectingRegion id="49C7F8272854FFC3F5C0FA4DFCC2B5DC" box="[679,865,1481,1506]" country="United States of America" name="New Hampshire" pageId="20" pageNumber="21">
New
<collectingRegion id="49C7F8272854FFC3F584FA4EFCC2B5DC" box="[739,865,1481,1506]" country="United Kingdom" name="Hampshire" pageId="20" pageNumber="21">Hampshire</collectingRegion>
</collectingRegion>
,
<collectingCountry id="F31476552854FFC3F40BFA4DFC07B5DC" box="[876,932,1482,1506]" name="United States of America" pageId="20" pageNumber="21">USA</collectingCountry>
(
<typeStatus id="54B888672854FFC3F4D4FA4EFBB9B5DC" box="[947,1050,1481,1506]" pageId="20" pageNumber="21" type="holotype">holotype</typeStatus>
, 1).
</materialsCitation>
</paragraph>
<paragraph id="8BBC36C52854FFC3F7A0FA6AFB20B638" blockId="20.[151,1437,1409,2010]" box="[199,1155,1517,1542]" pageId="20" pageNumber="21">
Published figures:
<bibRefCitation id="EF924B342854FFC3F6FEFA6AFCE2B638" author="Durden" box="[409,833,1517,1542]" pageId="20" pageNumber="21" refString="Durden, C. J. &amp; Rose, H. (1978) Butterflies from the Middle Miocene: the earliest occurrence of fossil Papilionoidea (Lepidoptera). Pearce-Sellards Series, 29, 1 - 25." type="journal article" year="1978">
Durden &amp; Rose (1978:
<figureCitation id="13382A402854FFC3F5C7FA6AFD46B638" box="[672,741,1517,1542]" 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="20" pageNumber="21">Figs 1</figureCitation>
, 6A–B)
</bibRefCitation>
;
<bibRefCitation id="EF924B342854FFC3F429FA69FBDCB638" author="Murata" box="[846,1151,1517,1542]" pageId="20" pageNumber="21" refString="Murata, Y. (1998) Notes on the evolutionary process of butterflies and butterfly fossils. Butterflies, 20 (part 1 &amp; 2), 4 - 17 &amp; 27 - 40. [in Japanese with English summary]" type="journal article" year="1998">
Murata (1998:
<figureCitation id="13382A402854FFC3F491FA6AFB98B638" box="[1014,1083,1517,1542]" 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="20" pageNumber="21">Figs 1</figureCitation>
,
<figureCitation id="13382A402854FFC3F32EFA69FBFAB638" box="[1097,1113,1518,1542]" 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="20" pageNumber="21">2</figureCitation>
,
<figureCitation id="13382A402854FFC3F302FA6AFBD6B638" box="[1125,1141,1517,1542]" 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="20" pageNumber="21">5</figureCitation>
)
</bibRefCitation>
.
</paragraph>
<paragraph id="8BBC36C52854FFC2F7A0F996FD13B193" blockId="20.[151,1437,1409,2010]" lastBlockId="21.[151,1437,151,1725]" lastPageId="21" lastPageNumber="22" pageId="20" pageNumber="21">
Rather well preserved, seen from above, with wings spread. Length of forewing
<quantity id="4CFB9B202854FFC3F3E0F995FB7DB617" box="[1159,1246,1554,1578]" metricMagnitude="-2" metricUnit="m" metricValue="3.7" pageId="20" pageNumber="21" unit="mm" value="37.0">37 mm</quantity>
. Two papilionid autapomorphies are present, both in the forewing: crossvein Cu-1A (Durden &amp; Rose: Cu-V;
<bibRefCitation id="EF924B342854FFC3F3AEF9B2FAF6B670" author="Miller" box="[1225,1365,1589,1614]" pageId="20" pageNumber="21" refString="Miller, J. S. (1987) Phylogenetic s tudies in the Papilioninae (Lepidoptera: Papilionidae). Bulletin American Museum Natural History, 186, 365 - 512." type="journal article" year="1987">Miller 1987</bibRefCitation>
: basal spur) present, and 2A (Durden &amp; Rose:
<date id="FFBD10052854FFC3F50DF9DDFD2DB64C" box="[618,654,1626,1650]" pageId="20" pageNumber="21">V3</date>
) terminates on dorsum. In the hindwing there are two anal veins, a primitive condition, which in the
<taxonomicName id="4C034D462854FFC3F573F9FAFD03B6A8" box="[532,672,1661,1686]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="20" pageNumber="21" phylum="Arthropoda" rank="family">Papilionidae</taxonomicName>
is found in
<taxonomicName id="4C034D462854FFC3F44CF9F9FBBFB6A8" authority="(Baroniinae)" baseAuthorityName="Baroniinae" box="[811,1052,1661,1686]" class="Insecta" family="Papilionidae" genus="Baronia" kingdom="Animalia" order="Lepidoptera" pageId="20" pageNumber="21" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72854FFC3F44CF9F9FC25B6AB" box="[811,902,1662,1685]" italics="true" pageId="20" pageNumber="21">Baronia</emphasis>
(Baroniinae)
</taxonomicName>
only. The posterior margin of the forewing cell is strongly trifid, a plesiomorphic condition not found in extant
<taxonomicName id="4C034D462854FFC3F371F926FB05B684" box="[1046,1190,1697,1722]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="20" pageNumber="21" phylum="Arthropoda" rank="family">Papilionidae</taxonomicName>
. In the forewing CuP seems to be present (interpreted by Durden &amp; Rose as a sclerotized crease), a primitive character not found in other Papilionoidea, although the course of the lost vein can be visible in extant
<taxonomicName id="4C034D462854FFC3F49BF96EFB21B73C" box="[1020,1154,1769,1794]" class="Insecta" family="Hesperiidae" kingdom="Animalia" order="Lepidoptera" pageId="20" pageNumber="21" phylum="Arthropoda" rank="family">Hesperiidae</taxonomicName>
by separate spots in the upper and lower part of the area between CuA2 and 1A+2A. Durden &amp; Rose also interpreted a possible crease in the hindwing as CuP, but this seems unlikely, since it seems to originate from the base of 1A+2A, and it may be an artefact. Contrary to what is found in
<taxonomicName id="4C034D462854FFC3F558F8D2FD6DB750" box="[575,718,1877,1902]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="20" pageNumber="21" phylum="Arthropoda" rank="subFamily" subFamily="Papilioninae">Papilioninae</taxonomicName>
, there are only four radial branches in the forewing (formula: 1, 2, 3+5 (according to
<bibRefCitation id="EF924B342854FFC3F6E8F8FEFDB8B7AC" author="Miller" box="[399,539,1913,1938]" pageId="20" pageNumber="21" refString="Miller, J. S. (1987) Phylogenetic s tudies in the Papilioninae (Lepidoptera: Papilionidae). Bulletin American Museum Natural History, 186, 365 - 512." type="journal article" year="1987">Miller 1987</bibRefCitation>
, R4 is missing); R2 originates at upper corner of cell.
<taxonomicName id="4C034D462854FFC3F3C6F8FDFB5FB7AF" box="[1185,1276,1914,1937]" class="Insecta" family="Papilionidae" genus="Baronia" kingdom="Animalia" order="Lepidoptera" pageId="20" pageNumber="21" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72854FFC3F3C6F8FDFB5FB7AF" box="[1185,1276,1914,1937]" italics="true" pageId="20" pageNumber="21">Baronia</emphasis>
</taxonomicName>
also has a 4- branched radius in the forewing, but so do
<taxonomicName id="4C034D462854FFC3F5EDF819FCA5B78B" box="[650,774,1950,1973]" class="Insecta" family="Papilionidae" genus="Parnassius" kingdom="Animalia" order="Lepidoptera" pageId="20" pageNumber="21" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72854FFC3F5EDF819FCA5B78B" box="[650,774,1950,1973]" italics="true" pageId="20" pageNumber="21">Parnassius</emphasis>
</taxonomicName>
and
<taxonomicName id="4C034D462854FFC3F425F819FC41B78B" box="[834,994,1950,1973]" class="Insecta" family="Papilionidae" genus="Hypermnestra" kingdom="Animalia" order="Lepidoptera" pageId="20" pageNumber="21" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72854FFC3F425F819FC41B78B" box="[834,994,1950,1973]" italics="true" pageId="20" pageNumber="21">Hypermnestra</emphasis>
</taxonomicName>
, as well as species of
<taxonomicName id="4C034D462854FFC3F38EF81AFAE6B788" box="[1257,1349,1949,1974]" class="Insecta" family="Pieridae" kingdom="Animalia" order="Lepidoptera" pageId="20" pageNumber="21" phylum="Arthropoda" rank="family">Pieridae</taxonomicName>
(where even more radial veins may be lost),
<taxonomicName id="4C034D462854FFC3F550F846FD1AB7E4" box="[567,697,1985,2010]" class="Insecta" family="Lycaenidae" kingdom="Animalia" order="Lepidoptera" pageId="20" pageNumber="21" phylum="Arthropoda" rank="family">Lycaenidae</taxonomicName>
,
<taxonomicName id="4C034D462854FFC3F5A2F846FCE3B7E4" box="[709,832,1985,2010]" class="Insecta" family="Riodinidae" kingdom="Animalia" order="Lepidoptera" pageId="20" pageNumber="21" phylum="Arthropoda" rank="family">Riodinidae</taxonomicName>
and
<taxonomicName id="4C034D462854FFC3F41FF846FBB1B7E4" box="[888,1042,1985,2010]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="20" pageNumber="21" phylum="Arthropoda" rank="family">Nymphalidae</taxonomicName>
, suggesting that the loss of a radial branch occurred more than once in the evolution of the butterflies. Therefore, this character state in
<taxonomicName id="4C034D462855FFC2F271FF10FA3FB08E" box="[1302,1436,151,176]" class="Insecta" family="Papilionidae" genus="Praepapilio" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72855FFC2F271FF10FA3FB08E" box="[1302,1436,151,176]" italics="true" pageId="21" pageNumber="22">Praepapilio</emphasis>
</taxonomicName>
cannot be seen as a measure of close affinity with
<taxonomicName id="4C034D462855FFC2F5B5FF3AFC8EB0EA" box="[722,813,189,212]" class="Insecta" family="Papilionidae" genus="Baronia" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72855FFC2F5B5FF3AFC8EB0EA" box="[722,813,189,212]" italics="true" pageId="21" pageNumber="22">Baronia</emphasis>
</taxonomicName>
. According to
<bibRefCitation id="EF924B342855FFC2F4B0FF3BFBCCB0EA" author="Miller" box="[983,1135,188,213]" pageId="21" pageNumber="22" refString="Miller, J. S. (1987) Phylogenetic s tudies in the Papilioninae (Lepidoptera: Papilionidae). Bulletin American Museum Natural History, 186, 365 - 512." type="journal article" year="1987">Miller (1987)</bibRefCitation>
the crossvein Cu-1A is an apomorphy of the subfamily
<taxonomicName id="4C034D462855FFC2F680FF58FDD4B0C6" box="[487,631,223,248]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="subFamily" subFamily="Papilioninae">Papilioninae</taxonomicName>
.
<bibRefCitation id="EF924B342855FFC2F5E3FF58FC2DB0C6" author="Durden" box="[644,910,223,248]" pageId="21" pageNumber="22" refString="Durden, C. J. &amp; Rose, H. (1978) Butterflies from the Middle Miocene: the earliest occurrence of fossil Papilionoidea (Lepidoptera). Pearce-Sellards Series, 29, 1 - 25." type="journal article" year="1978">Durden &amp; Rose (1978)</bibRefCitation>
state that the patagia have a narrow elongate sclerotization. This is an apomorphy of
<taxonomicName id="4C034D462855FFC2F510FE83FD51B123" box="[631,754,260,285]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="subFamily" subFamily="Baroniinae">Baroniinae</taxonomicName>
(
<bibRefCitation id="EF924B342855FFC2F461FE83FC4EB122" author="Ackery" box="[774,1005,260,285]" pageId="21" pageNumber="22" 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="B977EAD72855FFC2F400FE82FC3DB123" box="[871,926,260,285]" italics="true" pageId="21" pageNumber="22">et al</emphasis>
. 1999
</bibRefCitation>
), but these authors indicate that the sclerotization is variable in butterflies, although generally not so much reduced as in
<taxonomicName id="4C034D462855FFC2F32FFEAEFB00B17E" box="[1096,1187,297,320]" class="Insecta" family="Papilionidae" genus="Baronia" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72855FFC2F32FFEAEFB00B17E" box="[1096,1187,297,320]" italics="true" pageId="21" pageNumber="22">Baronia</emphasis>
</taxonomicName>
. It is unlikely that this character is well visible in the fossil. It is even more unlikely that Durden &amp; Rose have actually seen a narrow sclerotic band uniting the cervical sclerites beneath the neck, as they state in the list of characters agreeing with the
<taxonomicName id="4C034D462855FFC2F7F0FE13FE85B193" box="[151,294,404,429]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="family">Papilionidae</taxonomicName>
, in a fossil that is seen from above.
</paragraph>
<paragraph id="8BBC36C52855FFC2F7A0FE30FDFBB25E" blockId="21.[151,1437,151,1725]" pageId="21" pageNumber="22">
The remark by
<bibRefCitation id="EF924B342855FFC2F619FE30FD0FB1EE" author="Durden" box="[382,684,439,464]" pageId="21" pageNumber="22" refString="Durden, C. J. &amp; Rose, H. (1978) Butterflies from the Middle Miocene: the earliest occurrence of fossil Papilionoidea (Lepidoptera). Pearce-Sellards Series, 29, 1 - 25." type="journal article" year="1978">Durden &amp; Rose (1978: 6)</bibRefCitation>
that &quot;Based on wing shape, abdominal dimensions preserved as deformation of matrix, and position of abdominal margin of hindwing, this specimens is a female&quot;, must be considered speculative, and their remark (
<bibRefCitation id="EF924B342855FFC2F51FFE78FC33B226" author="Durden" box="[632,912,511,536]" pageId="21" pageNumber="22" refString="Durden, C. J. &amp; Rose, H. (1978) Butterflies from the Middle Miocene: the earliest occurrence of fossil Papilionoidea (Lepidoptera). Pearce-Sellards Series, 29, 1 - 25." type="journal article" year="1978">Durden &amp; Rose 1978: 8</bibRefCitation>
), based on a single specimen, that &quot;Range of individual variation within sex, between sexes, and within species may be comparable to that found in the modern
<taxonomicName id="4C034D462855FFC2F7F0FDCEFED4B25E" box="[151,375,583,608]" class="Insecta" family="Papilionidae" genus="Baronia" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="species" species="brevicornis">
<emphasis id="B977EAD72855FFC2F7F0FDCEFED4B25E" box="[151,375,583,608]" italics="true" pageId="21" pageNumber="22">Baronia brevicornis</emphasis>
</taxonomicName>
…&quot; lacks evidence.
</paragraph>
<paragraph id="8BBC36C52855FFC2F7A0FDEBFEA7B306" blockId="21.[151,1437,151,1725]" pageId="21" pageNumber="22">
Miller's (1987) remark that &quot;the discovery of
<taxonomicName id="4C034D462855FFC2F5A9FDEBFCF7B2BB" box="[718,852,620,645]" class="Insecta" family="Papilionidae" genus="Praepapilio" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72855FFC2F5A9FDEBFCF7B2BB" box="[718,852,620,645]" italics="true" pageId="21" pageNumber="22">Praepapilio</emphasis>
</taxonomicName>
confirms that genera within the
<taxonomicName id="4C034D462855FFC2F3AEFDEBFAF6B2BB" box="[1225,1365,620,645]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="family">Papilionidae</taxonomicName>
are at least 48 million years old&quot;, is not particularly meaningful, since the recognition of a supraspecific taxon is subjective and all extant genera could well be younger. Apart from
<taxonomicName id="4C034D462855FFC2F4D0FD33FB9EB2F3" box="[951,1085,692,717]" class="Insecta" family="Papilionidae" genus="Praepapilio" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72855FFC2F4D0FD33FB9EB2F3" box="[951,1085,692,717]" italics="true" pageId="21" pageNumber="22">Praepapilio</emphasis>
</taxonomicName>
many other genera may have existed that presently no longer occur, so why would extant genera be of such antiquity? The discovery of the fossil does confirm that some of the apomorphies of the
<taxonomicName id="4C034D462855FFC2F5A9FD7BFCFEB32B" box="[718,861,764,789]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="family">Papilionidae</taxonomicName>
,
<taxonomicName id="4C034D462855FFC2F40FFD7BFC40B32B" box="[872,995,764,789]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="subFamily" subFamily="Baroniinae">Baroniinae</taxonomicName>
and
<taxonomicName id="4C034D462855FFC2F37DFD7BFB05B32B" box="[1050,1190,764,789]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="subFamily" subFamily="Papilioninae">Papilioninae</taxonomicName>
are at least the age of the fossil.
</paragraph>
<paragraph id="8BBC36C52855FFC2F7A0FCC3FA3FB56A" blockId="21.[151,1437,151,1725]" pageId="21" pageNumber="22">
The genus
<taxonomicName id="4C034D462855FFC2F624FCC3FE6AB363" box="[323,457,836,861]" class="Insecta" family="Papilionidae" genus="Praepapilio" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72855FFC2F624FCC3FE6AB363" box="[323,457,836,861]" italics="true" pageId="21" pageNumber="22">Praepapilio</emphasis>
</taxonomicName>
and the subfamily
<taxonomicName id="4C034D462855FFC2F5C5FCC3FCC2B363" box="[674,865,836,861]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="subFamily" subFamily="Praepapilioninae">Praepapilioninae</taxonomicName>
, erected by
<bibRefCitation id="EF924B342855FFC2F48FFCC3FB48B362" author="Durden" box="[1000,1259,836,861]" pageId="21" pageNumber="22" refString="Durden, C. J. &amp; Rose, H. (1978) Butterflies from the Middle Miocene: the earliest occurrence of fossil Papilionoidea (Lepidoptera). Pearce-Sellards Series, 29, 1 - 25." type="journal article" year="1978">Durden &amp; Rose (1978)</bibRefCitation>
for this species and its supposed fossil congener
<taxonomicName id="4C034D462855FFC2F562FCEEFDD5B3BE" box="[517,630,871,896]" class="Insecta" family="Papilionidae" genus="Praepapilio" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="species" species="gracilis">
<emphasis id="B977EAD72855FFC2F562FCEEFDD5B3BE" box="[517,630,871,896]" italics="true" pageId="21" pageNumber="22">P. gracilis</emphasis>
</taxonomicName>
(see below), are not based on autapomorphic character states, but on the absence of apomorphies of other
<taxonomicName id="4C034D462855FFC2F56BFC0BFD3FB39B" box="[524,668,908,933]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="family">Papilionidae</taxonomicName>
. In recent analyses (e.g.
<bibRefCitation id="EF924B342855FFC2F4C8FC0BFB38B39A" author="Zakharov" box="[943,1179,908,933]" pageId="21" pageNumber="22" refString="Zakharov, E. V., Caterino, M. S. &amp; 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="B977EAD72855FFC2F346FC0AFBF2B39B" box="[1057,1105,908,933]" italics="true" pageId="21" pageNumber="22">et al</emphasis>
. 2004
</bibRefCitation>
;
<bibRefCitation id="EF924B342855FFC2F3C0FC0BFA36B39A" author="Simonsen" box="[1191,1429,908,933]" pageId="21" pageNumber="22" refString="Simonsen, T. J., Zakharov, E. V., Djernaes, M., Cotton, A. M., Vane-Wright, R. I. &amp; 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="B977EAD72855FFC2F27BFC0AFAEDB39B" box="[1308,1358,908,933]" italics="true" pageId="21" pageNumber="22">et al</emphasis>
. 2011
</bibRefCitation>
) the extant
<taxonomicName id="4C034D462855FFC2F672FC28FE02B3F6" box="[277,417,943,968]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="family">Papilionidae</taxonomicName>
are subdivided in
<taxonomicName id="4C034D462855FFC2F5E4FC28FCA2B3F6" box="[643,769,943,968]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="subFamily" subFamily="Baroniinae">Baroniinae</taxonomicName>
,
<taxonomicName id="4C034D462855FFC2F477FC28FC3FB3F6" box="[784,924,943,968]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="subFamily" subFamily="Parnassiinae">Parnassiinae</taxonomicName>
and
<taxonomicName id="4C034D462855FFC2F4BAFC28FBCFB3F6" box="[989,1132,943,968]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="subFamily" subFamily="Papilioninae">Papilioninae</taxonomicName>
, which are interrelated as follows:
<taxonomicName id="4C034D462855FFC2F667FC53FEDEB3D3" box="[256,381,980,1005]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="subFamily" subFamily="Baroniinae">Baroniinae</taxonomicName>
+(
<taxonomicName id="4C034D462855FFC2F6F4FC53FD81B3D3" box="[403,546,980,1005]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="subFamily" subFamily="Parnassiinae">Parnassiinae</taxonomicName>
+
<taxonomicName id="4C034D462855FFC2F556FC53FD62B3D3" box="[561,705,980,1005]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="subFamily" subFamily="Papilioninae">Papilioninae</taxonomicName>
). (The
<taxonomicName id="4C034D462855FFC2F47CFC53FC35B3D3" box="[795,918,980,1005]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="subFamily" subFamily="Baroniinae">Baroniinae</taxonomicName>
is monotypic, consisting solely of
<taxonomicName id="4C034D462855FFC2F226FC52FE1BB42E" authority="Salvin, 1893" authorityName="Salvin" authorityYear="1893" class="Insecta" family="Papilionidae" genus="Baronia" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="species" species="brevicornis">
<emphasis id="B977EAD72855FFC2F226FC52FEB6B42E" italics="true" pageId="21" pageNumber="22">Baronia brevicornis</emphasis>
Salvin, 1893
</taxonomicName>
.) An attempt to find the taxonomic position of
<taxonomicName id="4C034D462855FFC2F494FC70FBDAB42E" box="[1011,1145,1015,1040]" class="Insecta" family="Papilionidae" genus="Praepapilio" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72855FFC2F494FC70FBDAB42E" box="[1011,1145,1015,1040]" italics="true" pageId="21" pageNumber="22">Praepapilio</emphasis>
</taxonomicName>
in the phylogeny of the
<taxonomicName id="4C034D462855FFC2F7F0FB9BFE80B40B" box="[151,291,1052,1077]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="family">Papilionidae</taxonomicName>
is hampered by the fact that only venational characters of the fossil are helpful. On the basis of the trifid condition of the posterior margin of the forewing cell
<taxonomicName id="4C034D462855FFC2F41FFBB8FC5DB466" box="[888,1022,1087,1112]" class="Insecta" family="Papilionidae" genus="Praepapilio" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72855FFC2F41FFBB8FC5DB466" box="[888,1022,1087,1112]" italics="true" pageId="21" pageNumber="22">Praepapilio</emphasis>
</taxonomicName>
is more primitive than the extant
<taxonomicName id="4C034D462855FFC2F7F0FBE3FE84B443" box="[151,295,1124,1149]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="family">Papilionidae</taxonomicName>
. The primitive condition agrees with the apparent presence of (a remnant of) CuP in the forewing. Moreover, it shares with
<taxonomicName id="4C034D462855FFC2F6D6FB0EFDAFB49E" box="[433,524,1161,1184]" class="Insecta" family="Papilionidae" genus="Baronia" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72855FFC2F6D6FB0EFDAFB49E" box="[433,524,1161,1184]" italics="true" pageId="21" pageNumber="22">Baronia</emphasis>
</taxonomicName>
another plesiomorphic trait, the presence of two anal veins in the hindwing. The the basal spur in the forewing is a character only found in
<taxonomicName id="4C034D462855FFC2F45BFB2BFC68B4FB" box="[828,971,1196,1221]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="family">Papilionidae</taxonomicName>
, but it is absent in
<taxonomicName id="4C034D462855FFC2F3CEFB2BFA87B4FB" box="[1193,1316,1196,1221]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="subFamily" subFamily="Baroniinae">Baroniinae</taxonomicName>
and many
<taxonomicName id="4C034D462855FFC2F7F0FB48FE80B4D6" box="[151,291,1231,1256]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="subFamily" subFamily="Parnassiinae">Parnassiinae</taxonomicName>
as well as in the genus
<emphasis id="B977EAD72855FFC2F55AFB48FD66B4D6" box="[573,709,1231,1256]" italics="true" pageId="21" pageNumber="22">Teniopalpus</emphasis>
of the
<taxonomicName id="4C034D462855FFC2F47CFB48FC04B4D6" box="[795,935,1231,1256]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="subFamily" subFamily="Papilioninae">Papilioninae</taxonomicName>
(
<bibRefCitation id="EF924B342855FFC2F4DFFB48FB37B4D6" author="Ackery" box="[952,1172,1231,1256]" pageId="21" pageNumber="22" 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 et al. 1999</bibRefCitation>
), indicating that it was probably lost several times. Alternatively, it could quite well be a remnant of the otherwise lost CuP, and without further evidence its presence is no criterion for close relationship within
<taxonomicName id="4C034D462855FFC2F4DBFA90FBEFB50E" box="[956,1100,1303,1328]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="family">Papilionidae</taxonomicName>
. Summing up this evidence, if we want to use
<taxonomicName id="4C034D462855FFC2F622FABBFE68B56B" box="[325,459,1340,1365]" class="Insecta" family="Papilionidae" genus="Praepapilio" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72855FFC2F622FABBFE68B56B" box="[325,459,1340,1365]" italics="true" pageId="21" pageNumber="22">Praepapilio</emphasis>
</taxonomicName>
as a calibration point, it is best placed at the root of the
<taxonomicName id="4C034D462855FFC2F359FABBFB69B56B" box="[1086,1226,1340,1365]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="family">Papilionidae</taxonomicName>
and not higher up.
</paragraph>
</subSubSection>
<subSubSection id="C319654E2855FFC2F7A0FAD8FBC3B683" pageId="21" pageNumber="22" type="reference_group">
<paragraph id="8BBC36C52855FFC2F7A0FAD8FBC3B683" blockId="21.[151,1437,151,1725]" pageId="21" pageNumber="22">
Placing
<taxonomicName id="4C034D462855FFC2F645FAD8FE0BB546" box="[290,424,1375,1400]" class="Insecta" family="Papilionidae" genus="Praepapilio" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72855FFC2F645FAD8FE0BB546" box="[290,424,1375,1400]" italics="true" pageId="21" pageNumber="22">Praepapilio</emphasis>
</taxonomicName>
at the stem node of
<taxonomicName id="4C034D462855FFC2F5ECFAD8FCBAB546" box="[651,793,1375,1400]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="subFamily" subFamily="Parnassiinae">Parnassiinae</taxonomicName>
+
<taxonomicName id="4C034D462855FFC2F44FFAD8FC15B546" box="[808,950,1375,1400]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="subFamily" subFamily="Papilioninae">Papilioninae</taxonomicName>
,
<bibRefCitation id="EF924B342855FFC2F4A6FAD8FB1CB546" box="[961,1215,1375,1400]" pageId="21" pageNumber="22" refString="Simonsen, T. J., Zakharov, E. V., Djernaes, M., Cotton, A. M., Vane-Wright, R. I. &amp; 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">
Simonsen
<emphasis id="B977EAD72855FFC2F350FAE6FBCCB546" box="[1079,1135,1375,1400]" italics="true" pageId="21" pageNumber="22">et al.</emphasis>
(2011)
</bibRefCitation>
found an estimated time of origin of the
<taxonomicName id="4C034D462855FFC2F6EFFA03FDB7B5A3" ID-CoL="DX5" box="[392,532,1412,1437]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="family">Papilionidae</taxonomicName>
at 68 (53–87) Ma.
<bibRefCitation id="EF924B342855FFC2F593FA03FC76B5A2" box="[756,981,1412,1437]" pageId="21" pageNumber="22" refString="Nazari, V., Zakharov, E. V. &amp; 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="B977EAD72855FFC2F42FFA02FCD8B5A3" box="[840,891,1412,1437]" italics="true" pageId="21" pageNumber="22">et al</emphasis>
. (2007)
</bibRefCitation>
included the fossil as well as two other fossil
<taxonomicName id="4C034D462855FFC2F7BDFA20FECAB5FE" ID-CoL="DX5" box="[218,361,1447,1472]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="family">Papilionidae</taxonomicName>
, in their analysis of extant
<taxonomicName id="4C034D462855FFC2F5F2FA20FC87B5FE" ID-CoL="DX5" box="[661,804,1447,1472]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="family">Papilionidae</taxonomicName>
, making use of morphological and molecular characters, even though the fossils obviously lacked most characters. In this analysis
<taxonomicName id="4C034D462855FFC2F489FA4BFBD7B5DB" box="[1006,1140,1484,1509]" class="Insecta" family="Papilionidae" genus="Praepapilio" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72855FFC2F489FA4BFBD7B5DB" box="[1006,1140,1484,1509]" italics="true" pageId="21" pageNumber="22">Praepapilio</emphasis>
</taxonomicName>
turned up as sister to the genus
<taxonomicName id="4C034D462855FFC2F786FA68FE90B636" ID-CoL="6DFM" box="[225,307,1519,1544]" class="Insecta" family="Papilionidae" genus="Papilio" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72855FFC2F786FA68FE90B636" box="[225,307,1519,1544]" italics="true" pageId="21" pageNumber="22">Papilio</emphasis>
</taxonomicName>
. Using the fossils as well as some other information for calibration,
<bibRefCitation id="EF924B342855FFC2F32BFA68FA8DB636" box="[1100,1326,1519,1544]" pageId="21" pageNumber="22" refString="Nazari, V., Zakharov, E. V. &amp; 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="B977EAD72855FFC2F3C7FA76FB70B636" box="[1184,1235,1519,1544]" italics="true" pageId="21" pageNumber="22">et al</emphasis>
. (2007)
</bibRefCitation>
found an estimated age for the
<taxonomicName id="4C034D462855FFC2F6EDF993FDB5B613" ID-CoL="DX5" box="[394,534,1556,1581]" class="Insecta" family="Papilionidae" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="family">Papilionidae</taxonomicName>
of c. 100 million years, strongly contrasting with the age as found by
<bibRefCitation id="EF924B342855FFC2F249F993FE80B66E" pageId="21" pageNumber="22" refString="Simonsen, T. J., Zakharov, E. V., Djernaes, M., Cotton, A. M., Vane-Wright, R. I. &amp; 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">
Simonsen
<emphasis id="B977EAD72855FFC2F7F0F9BEFF69B66E" box="[151,202,1591,1616]" italics="true" pageId="21" pageNumber="22">et al</emphasis>
. (2011)
</bibRefCitation>
on the basis of a more realistic position of
<taxonomicName id="4C034D462855FFC2F47FF9B0FC3DB66E" box="[792,926,1591,1616]" class="Insecta" family="Papilionidae" genus="Praepapilio" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72855FFC2F47FF9B0FC3DB66E" box="[792,926,1591,1616]" italics="true" pageId="21" pageNumber="22">Praepapilio</emphasis>
</taxonomicName>
. The reason for this large difference is aptly discussed by
<bibRefCitation id="EF924B342855FFC2F648F9DBFD96B64A" box="[303,565,1628,1653]" pageId="21" pageNumber="22" refString="Simonsen, T. J., Zakharov, E. V., Djernaes, M., Cotton, A. M., Vane-Wright, R. I. &amp; 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">
Simonsen
<emphasis id="B977EAD72855FFC2F6C1F9DAFE43B64B" box="[422,480,1628,1653]" italics="true" pageId="21" pageNumber="22">et al.</emphasis>
(2011)
</bibRefCitation>
. Using
<taxonomicName id="4C034D462855FFC2F5EAF9DBFCB0B64B" box="[653,787,1628,1653]" class="Insecta" family="Papilionidae" genus="Praepapilio" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72855FFC2F5EAF9DBFCB0B64B" box="[653,787,1628,1653]" italics="true" pageId="21" pageNumber="22">Praepapilio</emphasis>
</taxonomicName>
as well as
<taxonomicName id="4C034D462855FFC2F4F3F9DBFBC7B64A" box="[916,1124,1628,1653]" class="Insecta" family="Papilionidae" genus="Thaites" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="species" species="ruminiana">
<emphasis id="B977EAD72855FFC2F4F3F9DBFBC7B64A" box="[916,1124,1628,1653]" italics="true" pageId="21" pageNumber="22">Thaites ruminiana</emphasis>
</taxonomicName>
and
<taxonomicName id="4C034D462855FFC2F3FAF9DAFAD0B64B" box="[1181,1395,1628,1653]" class="Insecta" family="Papilionidae" genus="Doritites" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="species" species="bosnaskii">
<emphasis id="B977EAD72855FFC2F3FAF9DAFAD0B64B" box="[1181,1395,1628,1653]" italics="true" pageId="21" pageNumber="22">Doritites bosnaskii</emphasis>
</taxonomicName>
for calibration,
<bibRefCitation id="EF924B342855FFC2F67AF9F8FD92B6A6" box="[285,561,1663,1688]" pageId="21" pageNumber="22" refString="Condamine, F. L., Sperling, F. A. H., Wahlberg, N., Rasplus, J. - Y. &amp; 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="B977EAD72855FFC2F6C1F906FE7BB6A6" box="[422,472,1663,1688]" italics="true" pageId="21" pageNumber="22">et al</emphasis>
. (2012)
</bibRefCitation>
estimated the earliest split in the papilionid lineage at about 52 Ma, agreeing with an age for the family not very different from Simonsen’s
<emphasis id="B977EAD72855FFC2F437F922FC22B683" box="[848,897,1700,1725]" italics="true" pageId="21" pageNumber="22">et al</emphasis>
. (2011) conclusion.
</paragraph>
</subSubSection>
</treatment>
</document>
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