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<mods:title id="0AF8DF2FDE260F6E6FE62A557421940F">Fossil butterflies, calibration points and the molecular clock (Lepidoptera: Papilionoidea)</mods:title>
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<treatment id="03AA87D3285BFFCBF7F0FB2CFF4CB15A" ID-DOI="http://doi.org/10.5281/zenodo.6046979" ID-GBIF-Taxon="130844917" ID-Zenodo-Dep="6046979" LSID="urn:lsid:plazi:treatment:03AA87D3285BFFCBF7F0FB2CFF4CB15A" httpUri="http://treatment.plazi.org/id/03AA87D3285BFFCBF7F0FB2CFF4CB15A" lastPageId="28" lastPageNumber="28" pageId="27" pageNumber="28">
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<paragraph id="8BBC36C5285BFFCCF7F0FB2CFF71B4D9" blockId="27.[151,1382,1195,1255]" pageId="27" pageNumber="28">
<heading id="D0F481A9285BFFCCF7F0FB2CFF71B4D9" bold="true" fontSize="11" level="1" pageId="27" pageNumber="28" reason="1">
<emphasis id="B977EAD7285BFFCCF7F0FB2CFF71B4D9" bold="true" pageId="27" pageNumber="28">
<emphasis id="B977EAD7285BFFCCF7F0FB2CFF54B4FB" bold="true" box="[151,247,1195,1221]" italics="true" pageId="27" pageNumber="28">
<taxonomicName id="4C034D46285BFFCCF7F0FB2CFF57B4FB" box="[151,244,1195,1221]" class="Insecta" family="Nymphalidae" genus="Neorinella" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="species" species="garciae">garciae</taxonomicName>
.
</emphasis>
<taxonomicName id="4C034D46285BFFCCF798FB2CFF71B4D9" authority="Martins-Neto, Kucera-Santos, de Moraes Vieira &amp; de Campos Fragoso, 1993" authorityName="Martins-Neto, Kucera-Santos, de Moraes Vieira &amp; de Campos Fragoso" authorityYear="1993" class="Insecta" family="Nymphalidae" genus="Neorinella" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="species" species="garciae">
<emphasis id="B977EAD7285BFFCCF798FB2CFE43B4FB" bold="true" box="[255,480,1195,1221]" italics="true" pageId="27" pageNumber="28">Neorinella garciae</emphasis>
Martins-Neto, Kucera-Santos, de Moraes Vieira &amp; de Campos Fragoso, 1993
</taxonomicName>
</emphasis>
</heading>
</paragraph>
<paragraph id="8BBC36C5285BFFCCF7F0FA93FD8FB513" blockId="27.[151,1437,1300,2009]" box="[151,556,1300,1325]" pageId="27" pageNumber="28">
<taxonomicName id="4C034D46285BFFCCF7F0FA93FE90B513" box="[151,307,1300,1325]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="family">Nymphalidae</taxonomicName>
:
<taxonomicName id="4C034D46285BFFCCF65AFA93FE09B513" box="[317,426,1300,1325]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="subFamily" subFamily="Satyrinae">Satyrinae</taxonomicName>
,
<taxonomicName id="4C034D46285BFFCCF6D3FA93FD8BB513" box="[436,552,1300,1325]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="tribe" tribe="Elymniini">Elymniini</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection id="C319654E285BFFCBF7A0FAB0FF4CB15A" lastPageId="28" lastPageNumber="29" pageId="27" pageNumber="28" type="description">
<paragraph id="8BBC36C5285BFFCCF7A0FAB0FEA3B54B" blockId="27.[151,1437,1300,2009]" pageId="27" pageNumber="28">
<materialsCitation id="3B6B3C98285BFFCCF7A0FAB0FEA3B54B" ID-GBIF-Occurrence="1503739224" country="Brazil" location="Sao Paulo" pageId="27" pageNumber="28" specimenCount="1" stateProvince="Sao Paulo">
<collectingCountry id="F3147655285BFFCCF7A0FAB0FEADB56E" box="[199,270,1335,1360]" name="Brazil" pageId="27" pageNumber="28">Brazil</collectingCountry>
, State of
<collectingRegion id="49C7F827285BFFCCF61EFABFFE4DB56E" box="[377,494,1335,1360]" country="Brazil" name="Sao Paulo" pageId="27" pageNumber="28">São Paulo</collectingRegion>
, Taubaté Basin, Tremembé Formation; ChattianAquitanian, late Oligocene—early Miocene.
</materialsCitation>
</paragraph>
<paragraph id="8BBC36C5285BFFCCF7A0FAF8FD85B5A6" blockId="27.[151,1437,1300,2009]" box="[199,550,1407,1432]" pageId="27" pageNumber="28">Depository: DGUG (holotype).</paragraph>
<paragraph id="8BBC36C5285BFFCCF7A0FA23FB67B582" blockId="27.[151,1437,1300,2009]" box="[199,1220,1444,1469]" pageId="27" pageNumber="28">
Published figures: Martins-Neto
<emphasis id="B977EAD7285BFFCCF551FA22FDC4B583" box="[566,615,1444,1469]" italics="true" pageId="27" pageNumber="28">et al</emphasis>
. (1993:
<figureCitation id="13382A40285BFFCCF5A1FA23FCAEB582" box="[710,781,1444,1469]" 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="27" pageNumber="28">Figs 2</figureCitation>
,
<figureCitation id="13382A40285BFFCCF47EFA23FC8BB582" box="[793,808,1444,1468]" captionStart="FIGURES 3 4" captionStartId="10.[151,264,1801,1823]" captionTargetBox="[165,1429,840,1774]" captionTargetId="figure@10.[154,1438,828,1788]" captionTargetPageId="10" captionText="FIGURES 3 4. Wing venation of (A) Pareronia Bingham, 1907 (Pieridae), and (B) Hasora Moore, 1881 (Hesperiidae). See text for explanation." httpUri="https://zenodo.org/record/583186/files/figure.png" pageId="27" pageNumber="28">3</figureCitation>
, pls I, II);
<bibRefCitation id="EF924B34285BFFCCF4F8FA22FB1CB582" author="Murata" box="[927,1215,1444,1469]" pageId="27" pageNumber="28" 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: Figs 911)</bibRefCitation>
.
</paragraph>
<paragraph id="8BBC36C5285BFFCCF7A0FA40FBC7B776" blockId="27.[151,1437,1300,2009]" pageId="27" pageNumber="28">
Partly visible forewings and hindwings and compressed body. In addition to wing shape and venation (partly visible) a paler colored band across forewings and hindwings is visible; there is an eyespot in the cell of the forewing, in R4-R5 (visible in the photo, not represented in the drawing), and in CuA1-CuA2 (?) of the forewing and, vaguely, in Cu1-Cu2 of the hindwing. The length of the forewing is
<quantity id="4CFB9B20285BFFCCF48FF9B3FBF2B672" box="[1000,1105,1588,1612]" metricMagnitude="-2" metricUnit="m" metricValue="3.17" pageId="27" pageNumber="28" unit="mm" value="31.7">31.7 mm</quantity>
.
<collectingRegion id="49C7F827285BFFCCF338F9B2FB22B672" box="[1119,1153,1589,1612]" country="Cameroon" name="North" pageId="27" pageNumber="28">No</collectingRegion>
visible swollen bases of forewing veins. Most remarkable in the venation of the forewing is the common origin of M3 and Cu1 from the lower distal corner of the cell. Among extant Papilionoidea it is a unique autapomorphy of the satyrine genus
<taxonomicName id="4C034D46285BFFCCF7F0F918FE02B686" authority="Miller 1968" authorityName="Miller" authorityYear="1968" box="[151,417,1695,1720]" class="Insecta" family="Nymphalidae" genus="Elymnias" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD7285BFFCCF7F0F918FF5CB686" box="[151,255,1695,1720]" italics="true" pageId="27" pageNumber="28">Elymnias</emphasis>
(
<bibRefCitation id="EF924B34285BFFCCF669F918FE3BB686" author="Miller" box="[270,408,1695,1720]" pageId="27" pageNumber="28" refString="Miller, L. D. (1968) The higher classification, phylogeny and zoogeography of the Satyridae (Lepidoptera). Memoirs American Entomological Society, 24, 1 - 174." type="journal article" year="1968">Miller 1968</bibRefCitation>
)
</taxonomicName>
. For the rest, the reconstruction of the wings (
<figureCitation id="13382A40285BFFCCF4C3F918FC44B686" box="[932,999,1695,1720]" 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="27" pageNumber="28">
Fig.
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</figureCitation>
in the original publication, reproduced here as Fig. 9) is hypothetical for those parts of the wings that are absent in the fossil. Moreover, in the forewing the radial branching cannot have been correctly reproduced: in
<figureCitation id="13382A40285BFFCCF432F960FC38B73E" box="[853,923,1767,1792]" 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="27" pageNumber="28">Fig. 2</figureCitation>
of the original description two radial veins branch off before the end of the cell, at which point R2 originates. One of these veins coalesces over some distance with R2 before ending on the front margin. This part of the wing is absent in the fossil.
</paragraph>
<paragraph id="8BBC36C5285BFFCBF7A0F8D3FF4CB15A" blockId="27.[151,1437,1300,2009]" lastBlockId="28.[151,1436,151,357]" lastPageId="28" lastPageNumber="29" pageId="27" pageNumber="28">
The fossil, which has been placed in a genus of its own, is supposed (in the original publication) to be related to the extant genus
<taxonomicName id="4C034D46285BFFCCF63FF8FEFE17B7AE" box="[344,436,1913,1936]" class="Insecta" family="Nymphalidae" genus="Neorina" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD7285BFFCCF63FF8FEFE17B7AE" box="[344,436,1913,1936]" italics="true" pageId="27" pageNumber="28">Neorina</emphasis>
</taxonomicName>
(
<taxonomicName id="4C034D46285BFFCCF6A4F8F0FDFCB7AE" box="[451,607,1911,1936]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="family">Nymphalidae</taxonomicName>
,
<taxonomicName id="4C034D46285BFFCCF50BF8F0FD74B7AE" box="[620,727,1911,1936]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="subFamily" subFamily="Satyrinae">Satyrinae</taxonomicName>
) as well as to the fossil genera
<taxonomicName id="4C034D46285BFFCCF358F8FEFB1FB7AE" box="[1087,1212,1913,1936]" class="Insecta" family="Nymphalidae" genus="Neorinopis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD7285BFFCCF358F8FEFB1FB7AE" box="[1087,1212,1913,1936]" italics="true" pageId="27" pageNumber="28">Neorinopis</emphasis>
</taxonomicName>
(see under
<taxonomicName id="4C034D46285BFFCCF225F8F0FA30B7AE" box="[1346,1427,1911,1936]" class="Insecta" family="Nymphalidae" genus="Cyllo" higherTaxonomySource="CoL" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="species" species="sepulta">
<emphasis id="B977EAD7285BFFCCF225F8F0FA30B7AE" box="[1346,1427,1911,1936]" italics="true" pageId="27" pageNumber="28">sepulta</emphasis>
</taxonomicName>
) and
<taxonomicName id="4C034D46285BFFCCF7AEF81BFED0B78B" box="[201,371,1948,1973]" class="Insecta" family="Nymphalidae" genus="Pseudoneorina" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD7285BFFCCF7AEF81BFED0B78B" box="[201,371,1948,1973]" italics="true" pageId="27" pageNumber="28">Pseudoneorina</emphasis>
</taxonomicName>
(see under
<taxonomicName id="4C034D46285BFFCCF69BF81BFDF3B78B" box="[508,592,1948,1973]" class="Insecta" family="Nymphalidae" genus="Pseudoneorina" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="species" species="coulleti">
<emphasis id="B977EAD7285BFFCCF69BF81BFDF3B78B" box="[508,592,1948,1973]" italics="true" pageId="27" pageNumber="28">coulleti</emphasis>
</taxonomicName>
). Apparently, this is based on similarity in wing shape and wing pattern. Realizing the variation in these characters for extant
<taxonomicName id="4C034D46285BFFCCF58BF846FCEBB7E6" box="[748,840,1985,2008]" class="Insecta" family="Nymphalidae" genus="Neorina" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD7285BFFCCF58BF846FCEBB7E6" box="[748,840,1985,2008]" italics="true" pageId="27" pageNumber="28">Neorina</emphasis>
</taxonomicName>
species as well as in related genera, these characters are unreliable indicators of relationship. A structural character like venation appears more reliable. In combination with
<emphasis id="B977EAD7285CFFCBF7B1FF3AFEFEB0EA" box="[214,349,189,212]" italics="true" pageId="28" pageNumber="29">Neorinopsis</emphasis>
and
<taxonomicName id="4C034D46285CFFCBF6C7FF3BFDE9B0EB" box="[416,586,188,213]" class="Insecta" family="Nymphalidae" genus="Pseudoneorina" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="28" pageNumber="29" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD7285CFFCBF6C7FF3BFDE9B0EB" box="[416,586,188,213]" italics="true" pageId="28" pageNumber="29">Pseudoneorina</emphasis>
</taxonomicName>
,
<taxonomicName id="4C034D46285CFFCBF539FF3BFD75B0EB" box="[606,726,188,213]" class="Insecta" family="Nymphalidae" genus="Neorinella" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="28" pageNumber="29" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD7285CFFCBF539FF3BFD75B0EB" box="[606,726,188,213]" italics="true" pageId="28" pageNumber="29">Neorinella</emphasis>
</taxonomicName>
has been classified, without comment, as belonging to the subtribe
<taxonomicName id="4C034D46285CFFCBF79DFF58FEECB0C6" box="[250,335,223,248]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="28" pageNumber="29" phylum="Arthropoda" rank="subTribe" subTribe="Lethina">Lethina</taxonomicName>
of the tribe
<taxonomicName id="4C034D46285CFFCBF686FF58FD9FB0C6" box="[481,572,223,248]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="28" pageNumber="29" phylum="Arthropoda" rank="tribe" tribe="Satyrini">Satyrini</taxonomicName>
, while
<taxonomicName id="4C034D46285CFFCBF5F5FF66FD4DB0C6" box="[658,750,225,248]" class="Insecta" family="Nymphalidae" genus="Neorina" kingdom="Animalia" order="Lepidoptera" pageId="28" pageNumber="29" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD7285CFFCBF5F5FF66FD4DB0C6" box="[658,750,225,248]" italics="true" pageId="28" pageNumber="29">Neorina</emphasis>
</taxonomicName>
has been placed in the tribe
<taxonomicName id="4C034D46285CFFCBF35AFF58FB07B0C6" box="[1085,1188,223,248]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="28" pageNumber="29" phylum="Arthropoda" rank="tribe" tribe="Zetherini">Zetherini</taxonomicName>
by
<bibRefCitation id="EF924B34285CFFCBF3B5FF58FF47B123" author="Wahlberg" pageId="28" pageNumber="29" 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>
. This, again, must be based on wing shape and design. If we take venation as indicator of relationship, both
<taxonomicName id="4C034D46285CFFCBF7F0FEA0FEACB17E" box="[151,271,295,320]" class="Insecta" family="Nymphalidae" genus="Neorinella" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="28" pageNumber="29" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD7285CFFCBF7F0FEA0FEACB17E" box="[151,271,295,320]" italics="true" pageId="28" pageNumber="29">Neorinella</emphasis>
</taxonomicName>
and
<emphasis id="B977EAD7285CFFCBF635FEAEFE79B17E" box="[338,474,297,320]" italics="true" pageId="28" pageNumber="29">Neorinopsis</emphasis>
should be classified as
<taxonomicName id="4C034D46285CFFCBF465FEA0FCD5B17E" box="[770,886,295,320]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="28" pageNumber="29" phylum="Arthropoda" rank="tribe" tribe="Elymniini">Elymniini</taxonomicName>
. See further the discussion under
<emphasis id="B977EAD7285CFFCBF273FEAEFF4BB15B" italics="true" pageId="28" pageNumber="29">
Neorinopsis
<taxonomicName id="4C034D46285CFFCBF7F0FECBFF4BB15B" box="[151,232,332,357]" class="Insecta" family="Nymphalidae" genus="Cyllo" higherTaxonomySource="CoL" kingdom="Animalia" order="Lepidoptera" pageId="28" pageNumber="29" phylum="Arthropoda" rank="species" species="sepulta">sepulta</taxonomicName>
</emphasis>
.
</paragraph>
</subSubSection>
</treatment>
</document>