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<document id="A2172EA5EF77268A1CCD797F9C6858CF" 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="03AA87D32865FFF1F7F0FD51FEE0B56A" 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="Cyllo sepulta" docType="treatment" docVersion="9" lastPageNumber="39" masterDocId="FF93FFAB2840FFD7F767FF87FFA3B03E" masterDocTitle="Fossil butterflies, calibration points and the molecular clock (Lepidoptera: Papilionoidea)" masterLastPageNumber="63" masterPageNumber="1" pageNumber="38" updateTime="1698439901205" updateUser="ExternalLinkService">
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<mods:title id="6B40037C2EB09044C6E22CC3EA5C542A">Fossil butterflies, calibration points and the molecular clock (Lepidoptera: Papilionoidea)</mods:title>
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<mods:namePart id="66C6ED9DD67C181CF25E4BB21F932721">Jong, Rienk De</mods:namePart>
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<treatment id="03AA87D32865FFF1F7F0FD51FEE0B56A" ID-DOI="http://doi.org/10.5281/zenodo.6047049" ID-GBIF-Taxon="130844840" ID-Zenodo-Dep="6047049" LSID="urn:lsid:plazi:treatment:03AA87D32865FFF1F7F0FD51FEE0B56A" httpUri="http://treatment.plazi.org/id/03AA87D32865FFF1F7F0FD51FEE0B56A" lastPageId="38" lastPageNumber="39" pageId="37" pageNumber="38">
<subSubSection id="C319654E2865FFF2F7F0FD51FF4EB32C" pageId="37" pageNumber="38" type="nomenclature">
<paragraph id="8BBC36C52865FFF2F7F0FD51FDC6B2CE" blockId="37.[151,613,726,786]" box="[151,613,726,752]" pageId="37" pageNumber="38">
<heading id="D0F481A92865FFF2F7F0FD51FDC6B2CE" bold="true" box="[151,613,726,752]" fontSize="11" level="1" pageId="37" pageNumber="38" reason="1">
<emphasis id="B977EAD72865FFF2F7F0FD51FDC6B2CE" bold="true" box="[151,613,726,752]" pageId="37" pageNumber="38">
<emphasis id="B977EAD72865FFF2F7F0FD51FF57B2CE" bold="true" box="[151,244,726,752]" italics="true" pageId="37" pageNumber="38">
<taxonomicName id="4C034D462865FFF2F7F0FD51FF53B2CE" box="[151,240,726,752]" class="Insecta" family="Nymphalidae" genus="Cyllo" higherTaxonomySource="CoL" kingdom="Animalia" order="Lepidoptera" pageId="37" pageNumber="38" phylum="Arthropoda" rank="species" species="sepulta">sepulta</taxonomicName>
.
</emphasis>
<taxonomicName id="4C034D462865FFF2F79CFD51FDC6B2CE" authority="Boisduval, 1841" authorityName="Boisduval" authorityYear="1841" box="[251,613,726,752]" class="Insecta" family="Nymphalidae" genus="Cyllo" higherTaxonomySource="CoL" kingdom="Animalia" order="Lepidoptera" pageId="37" pageNumber="38" phylum="Arthropoda" rank="species" species="sepulta">
<emphasis id="B977EAD72865FFF2F79CFD51FE3BB2CE" bold="true" box="[251,408,726,752]" italics="true" pageId="37" pageNumber="38">Cyllo sepulta</emphasis>
<bibRefCitation id="EF924B342865FFF2F6F8FD51FDC6B2CE" author="Boisduval" box="[415,613,726,752]" pageId="37" pageNumber="38" refString="Boisduval, J. A. (1841) [Cyllo sepulta]. Annales de la Societe entomologique de France, 9, 371 - 374." type="journal article" year="1841">Boisduval, 1841</bibRefCitation>
</taxonomicName>
</emphasis>
</heading>
</paragraph>
<paragraph id="8BBC36C52865FFF2F7F0FD7EFF4EB32C" blockId="37.[151,613,726,786]" box="[151,237,761,786]" pageId="37" pageNumber="38">Fig. 17.</paragraph>
</subSubSection>
<subSubSection id="C319654E2865FFF2F7F0FCC6FBF0B6E3" pageId="37" pageNumber="38" type="description">
<paragraph id="8BBC36C52865FFF2F7F0FCC6FE0DB364" blockId="37.[151,1437,833,2010]" box="[151,430,833,858]" pageId="37" pageNumber="38">
<taxonomicName id="4C034D462865FFF2F7F0FCC6FE90B364" box="[151,307,833,858]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="37" pageNumber="38" phylum="Arthropoda" rank="family">Nymphalidae</taxonomicName>
:
<taxonomicName id="4C034D462865FFF2F65AFCC6FE08B364" box="[317,427,833,858]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="37" pageNumber="38" phylum="Arthropoda" rank="subFamily" subFamily="Satyrinae">Satyrinae</taxonomicName>
.
</paragraph>
<paragraph id="8BBC36C52865FFF2F7A0FCE1FA23B340" blockId="37.[151,1437,833,2010]" box="[199,1408,869,894]" pageId="37" pageNumber="38">France, Bouches-du-Rhône, Aix-en-Provence, France; Chattian–Aquitanian, late Oligocene–early Miocene.</paragraph>
<paragraph id="8BBC36C52865FFF2F7A0FC0EFD77B39C" blockId="37.[151,1437,833,2010]" box="[199,724,905,930]" pageId="37" pageNumber="38">Depository: MNHN (holotype, IPM B-24309).</paragraph>
<paragraph id="8BBC36C52865FFF2F7A0FC2AFDFBB3D4" blockId="37.[151,1437,833,2010]" pageId="37" pageNumber="38">
Published figures:
<bibRefCitation id="EF924B342865FFF2F6FEFC29FD5EB3F8" author="Leestmans" box="[409,765,941,966]" pageId="37" pageNumber="38" 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 6–8, 18)</bibRefCitation>
;
<bibRefCitation id="EF924B342865FFF2F46DFC29FB98B3F8" author="Murata" box="[778,1083,941,966]" pageId="37" pageNumber="38" 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 17–23)</bibRefCitation>
;
<bibRefCitation id="EF924B342865FFF2F320FC2AFF47B3D4" author="Nel" pageId="37" pageNumber="38" refString="Nel, A. &amp; Nel, J. (1986) A propos de deux Lepidopteres fossiles du Stampien d'Aix- en-Provence (Lepidoptera, Hesperiidae et Nymphalidae Satyrinae), Bulletin du Museum National d' Histoire Naturelle, 8, 343 - 350." type="journal article" year="1986" yearSuffix="a">
Nel &amp; Nel (1986:
<figureCitation id="13382A402865FFF2F275FC2AFAF7B3F8" box="[1298,1364,941,966]" 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="37" pageNumber="38">Fig. 2</figureCitation>
a, Pl. I
<figureCitation id="13382A402865FFF2F7F0FC56FF79B3D4" box="[151,218,977,1002]" 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="37" pageNumber="38">Fig. 2</figureCitation>
)
</bibRefCitation>
; Scudder (1875: Pl. I Figs 8–17).
</paragraph>
<paragraph id="8BBC36C52865FFF2F7A0FC72FC4AB534" blockId="37.[151,1437,833,2010]" pageId="37" pageNumber="38">
Original description (
<bibRefCitation id="EF924B342865FFF2F6DEFC72FDCEB430" author="Boisduval" box="[441,621,1013,1038]" pageId="37" pageNumber="38" refString="Boisduval, J. A. (1841) [Cyllo sepulta]. Annales de la Societe entomologique de France, 9, 371 - 374." type="journal article" year="1841">Boisduval 1841</bibRefCitation>
) cited by Scudder (1875) who dedicated a long discussion to the fossil. Further discussed by
<bibRefCitation id="EF924B342865FFF2F6E3FB9EFDE8B40C" author="Nel" box="[388,587,1049,1074]" pageId="37" pageNumber="38" refString="Nel, A. &amp; Nel, J. (1986) A propos de deux Lepidopteres fossiles du Stampien d'Aix- en-Provence (Lepidoptera, Hesperiidae et Nymphalidae Satyrinae), Bulletin du Museum National d' Histoire Naturelle, 8, 343 - 350." type="journal article" year="1986">Nel &amp; Nel (1986)</bibRefCitation>
. The wing venation and pattern is rather well preserved. The thorax and legs (hindlegs only) lack diagnostic characters. The forewing radial formula is 1, 2+(3+(4+5)) according to Scudder, 1, 2, 3+(4+5) according to Nel &amp; Nel (who studied the
<typeStatus id="54B888672865FFF2F580FBE6FCEEB444" box="[743,845,1121,1146]" pageId="37" pageNumber="38" type="holotype">holotype</typeStatus>
), M3 and Cu1 originating from the same point at the lower distal end of the cell, meaning that ldc runs from the origin of Cu1 to M2, cell closed but crossvein weak, basally swollen veins absent. Hindwing with two anal veins, cell closed but crossvein weak. Forewing with falcate apex, hindwing forming a tail at M3. Scudder considered the visible wing pattern to be that of the upperside, but
<bibRefCitation id="EF924B342865FFF2F7F0FB76FEFEB534" author="Nel" box="[151,349,1265,1290]" pageId="37" pageNumber="38" refString="Nel, A. &amp; Nel, J. (1986) A propos de deux Lepidopteres fossiles du Stampien d'Aix- en-Provence (Lepidoptera, Hesperiidae et Nymphalidae Satyrinae), Bulletin du Museum National d' Histoire Naturelle, 8, 343 - 350." type="journal article" year="1986">Nel &amp; Nel (1986)</bibRefCitation>
convincingly showed it to be the pattern of the underside.
</paragraph>
<paragraph id="8BBC36C52865FFF2F7A0FA92FBF0B6E3" blockId="37.[151,1437,833,2010]" pageId="37" pageNumber="38">
This specimen is generally placed in
<taxonomicName id="4C034D462865FFF2F5E0FA92FD52B510" box="[647,753,1301,1326]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="37" pageNumber="38" phylum="Arthropoda" rank="subFamily" subFamily="Satyrinae">Satyrinae</taxonomicName>
on basis of wing shape and pattern (submarginal ocelli).
<bibRefCitation id="EF924B342865FFF2F7F0FABEFEFDB56C" author="Boisduval" box="[151,350,1337,1362]" pageId="37" pageNumber="38" refString="Boisduval, J. A. (1841) [Cyllo sepulta]. Annales de la Societe entomologique de France, 9, 371 - 374." type="journal article" year="1841">Boisduval (1841)</bibRefCitation>
assigned it to a recent genus,
<taxonomicName id="4C034D462865FFF2F5ACFABEFCABB56C" box="[715,776,1337,1362]" class="Insecta" family="Nymphalidae" genus="Cyllo" higherTaxonomySource="CoL" kingdom="Animalia" order="Lepidoptera" pageId="37" pageNumber="38" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72865FFF2F5ACFABEFCABB56C" box="[715,776,1337,1362]" italics="true" pageId="37" pageNumber="38">Cyllo</emphasis>
</taxonomicName>
, which is a junior synonym of
<taxonomicName id="4C034D462865FFF2F3E6FABEFB48B56C" box="[1153,1259,1337,1362]" class="Insecta" family="Nymphalidae" genus="Melanitis" kingdom="Animalia" order="Lepidoptera" pageId="37" pageNumber="38" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72865FFF2F3E6FABEFB48B56C" box="[1153,1259,1337,1362]" italics="true" pageId="37" pageNumber="38">Melanitis</emphasis>
</taxonomicName>
, see
<bibRefCitation id="EF924B342865FFF2F24BFABEFF47B548" author="Hemming" pageId="37" pageNumber="38" refString="Hemming, F. (1967) The generic names of the butterflies and their type-species (Lepidoptera: Rhopalocera). Bulletin of the British Museum (Natural History), Entomology, 9 (Supplement), 1 - 509." type="journal article" year="1967">Hemming (1967)</bibRefCitation>
.
<bibRefCitation id="EF924B342865FFF2F788FADAFEDDB548" author="Butler" box="[239,382,1373,1398]" pageId="37" pageNumber="38" refString="Butler, A. G. (1873) On fossil butterflies. Lepidoptera Exotica, 15, 126 - 128." type="journal article" year="1873">Butler (1873</bibRefCitation>
, cited by Scudder 1875) erected the genus
<emphasis id="B977EAD72865FFF2F43DFAD9FC42B54B" box="[858,993,1374,1397]" italics="true" pageId="37" pageNumber="38">Neorinopsis</emphasis>
, although Scudder misspelled the fossil
<taxonomicName id="4C034D462865FFF2F7F0FA05FEB0B5A7" box="[151,275,1410,1433]" class="Insecta" family="Nymphalidae" genus="Neorinopis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="37" pageNumber="38" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72865FFF2F7F0FA05FEB0B5A7" box="[151,275,1410,1433]" italics="true" pageId="37" pageNumber="38">Neorinopis</emphasis>
</taxonomicName>
, repeated by later authors. In wingspan and pattern it is intermediate between recent satyrine genera
<taxonomicName id="4C034D462865FFF2F7F0FA21FF50B583" box="[151,243,1446,1469]" class="Insecta" family="Nymphalidae" genus="Neorina" kingdom="Animalia" order="Lepidoptera" pageId="37" pageNumber="38" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72865FFF2F7F0FA21FF50B583" box="[151,243,1446,1469]" italics="true" pageId="37" pageNumber="38">Neorina</emphasis>
</taxonomicName>
(Southeast Asia),
<taxonomicName id="4C034D462865FFF2F6BBFA22FDF5B580" box="[476,598,1445,1470]" class="Insecta" family="Nymphalidae" genus="Antirrhaea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="37" pageNumber="38" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72865FFF2F6BBFA22FDF5B580" box="[476,598,1445,1470]" italics="true" pageId="37" pageNumber="38">Antirrhaea</emphasis>
</taxonomicName>
(
<collectingRegion id="49C7F8272865FFF2F508FA22FD11B580" box="[623,690,1445,1470]" country="Tanzania" name="Kusini Unguja" pageId="37" pageNumber="38">South</collectingRegion>
America), and
<emphasis id="B977EAD72865FFF2F41CFA22FBB3B580" box="[891,1040,1445,1470]" italics="true" pageId="37" pageNumber="38">Anchiphlebia</emphasis>
, a junior objective synonym of
<taxonomicName id="4C034D462865FFF2F7F0FA4EFEB1B5DC" box="[151,274,1481,1506]" class="Insecta" family="Nymphalidae" genus="Antirrhaea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="37" pageNumber="38" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72865FFF2F7F0FA4EFEB1B5DC" box="[151,274,1481,1506]" italics="true" pageId="37" pageNumber="38">Antirrhaea</emphasis>
</taxonomicName>
(see
<bibRefCitation id="EF924B342865FFF2F63EFA4EFDB1B5DC" author="Hemming" box="[345,530,1481,1506]" pageId="37" pageNumber="38" refString="Hemming, F. (1967) The generic names of the butterflies and their type-species (Lepidoptera: Rhopalocera). Bulletin of the British Museum (Natural History), Entomology, 9 (Supplement), 1 - 509." type="journal article" year="1967">Hemming 1967</bibRefCitation>
).
<bibRefCitation id="EF924B342865FFF2F54BFA4EFCA5B5DC" author="Nel" box="[556,774,1481,1506]" pageId="37" pageNumber="38" refString="Nel, A. &amp; Nel, J. (1986) A propos de deux Lepidopteres fossiles du Stampien d'Aix- en-Provence (Lepidoptera, Hesperiidae et Nymphalidae Satyrinae), Bulletin du Museum National d' Histoire Naturelle, 8, 343 - 350." type="journal article" year="1986">Nel &amp; Nel (1986)</bibRefCitation>
are clear in their opinion about this similarity: “… le rapprochement indiqué par Scudder (1875, tableau p. 28) avec des genres sud-américains (
<emphasis id="B977EAD72865FFF2F26BFA6AFA37B638" box="[1292,1428,1517,1542]" italics="true" pageId="37" pageNumber="38">Anthirrhaea</emphasis>
,
<emphasis id="B977EAD72865FFF2F7F0F996FE8FB614" box="[151,300,1553,1578]" italics="true" pageId="37" pageNumber="38">Anchiphlebia</emphasis>
), perd ainsi toute raison d’être.” (“… therefore, the similarity to
<collectingRegion id="49C7F8272865FFF2F31AF996FB1DB614" box="[1149,1214,1553,1578]" country="Tanzania" name="Kusini Unguja" pageId="37" pageNumber="38">South</collectingRegion>
American genera (
<emphasis id="B977EAD72865FFF2F7C7F9B2FE8BB670" box="[160,296,1589,1614]" italics="true" pageId="37" pageNumber="38">Anthirrhaea</emphasis>
,
<emphasis id="B977EAD72865FFF2F65BF9B2FE72B670" box="[316,465,1589,1614]" italics="true" pageId="37" pageNumber="38">Anchiphlebia</emphasis>
), indicated by Scudder (1875, table p. 28), lacks any credibility.”). According to Scudder (1875), the venation is reminiscent of
<taxonomicName id="4C034D462865FFF2F5D7F9DDFCAFB64F" box="[688,780,1626,1649]" class="Insecta" family="Nymphalidae" genus="Neorina" kingdom="Animalia" order="Lepidoptera" pageId="37" pageNumber="38" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72865FFF2F5D7F9DDFCAFB64F" box="[688,780,1626,1649]" italics="true" pageId="37" pageNumber="38">Neorina</emphasis>
</taxonomicName>
,
<taxonomicName id="4C034D462865FFF2F47BF9DEFCF9B64C" box="[796,858,1625,1650]" class="Insecta" family="Nymphalidae" genus="Lethe" kingdom="Animalia" order="Lepidoptera" pageId="37" pageNumber="38" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72865FFF2F47BF9DEFCF9B64C" box="[796,858,1625,1650]" italics="true" pageId="37" pageNumber="38">Lethe</emphasis>
</taxonomicName>
,
<taxonomicName id="4C034D462865FFF2F40DF9DEFC41B64C" box="[874,994,1625,1650]" class="Insecta" family="Nymphalidae" genus="Zophoessa" higherTaxonomySource="CoL" kingdom="Animalia" order="Lepidoptera" pageId="37" pageNumber="38" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72865FFF2F40DF9DEFC41B64C" box="[874,994,1625,1650]" italics="true" pageId="37" pageNumber="38">Zophoessa</emphasis>
</taxonomicName>
and
<taxonomicName id="4C034D462865FFF2F37BF9DEFBFEB64C" box="[1052,1117,1625,1650]" class="Insecta" family="Nymphalidae" genus="Debis" higherTaxonomySource="CoL" kingdom="Animalia" order="Lepidoptera" pageId="37" pageNumber="38" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72865FFF2F37BF9DEFBFEB64C" box="[1052,1117,1625,1650]" italics="true" pageId="37" pageNumber="38">Debis</emphasis>
</taxonomicName>
, of which the latter two are synonyms of
<taxonomicName id="4C034D462865FFF2F654F9FAFED1B6A8" box="[307,370,1661,1686]" class="Insecta" family="Nymphalidae" genus="Lethe" kingdom="Animalia" order="Lepidoptera" pageId="37" pageNumber="38" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72865FFF2F654F9FAFED1B6A8" box="[307,370,1661,1686]" italics="true" pageId="37" pageNumber="38">Lethe</emphasis>
</taxonomicName>
(see
<bibRefCitation id="EF924B342865FFF2F6D5F9FAFD9DB6A8" author="Gaede" box="[434,574,1661,1686]" pageId="37" pageNumber="38" refString="Gaede, M. (1931) Satyridae. Lepidopterorum Catalogus, 29, 1 - 759." type="journal article" year="1931">Gaede 1931</bibRefCitation>
), but the radial vein arrangement is different, a remarkable observation by Scudder, since the common origin of M3 and Cu1 of the forewing in
<emphasis id="B977EAD72865FFF2F4DBF925FBE0B687" box="[956,1091,1698,1721]" italics="true" pageId="37" pageNumber="38">Neorinopsis</emphasis>
does not agree at all with the situation in these genera, but is similar to what is found in the extant genus
<taxonomicName id="4C034D462865FFF2F484F942FBE8B6E0" box="[995,1099,1733,1758]" class="Insecta" family="Nymphalidae" genus="Elymnias" kingdom="Animalia" order="Lepidoptera" pageId="37" pageNumber="38" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72865FFF2F484F942FBE8B6E0" box="[995,1099,1733,1758]" italics="true" pageId="37" pageNumber="38">Elymnias</emphasis>
</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection id="C319654E2865FFF1F7A0F96EFEE0B56A" lastPageId="38" lastPageNumber="39" pageId="37" pageNumber="38" type="reference_group">
<paragraph id="8BBC36C52865FFF2F7A0F96EFD64B788" blockId="37.[151,1437,833,2010]" pageId="37" pageNumber="38">
The
<taxonomicName id="4C034D462865FFF2F79AF96EFEC4B73C" ID-CoL="93MZX" box="[253,359,1769,1794]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="37" pageNumber="38" phylum="Arthropoda" rank="subFamily" subFamily="Satyrinae">Satyrinae</taxonomicName>
do not have a unique and universal morphological apomorphy (
<bibRefCitation id="EF924B342865FFF2F33DF96EFA99B73C" author="Ackery" box="[1114,1338,1769,1794]" pageId="37" pageNumber="38" 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="B977EAD72865FFF2F3DFF96DFB4EB73C" box="[1208,1261,1769,1794]" italics="true" pageId="37" pageNumber="38">et al</emphasis>
. 1999
</bibRefCitation>
; but see
<bibRefCitation id="EF924B342865FFF2F7F0F88AFE00B718" author="Freitas" box="[151,419,1805,1830]" pageId="37" pageNumber="38" refString="Freitas, A. V. L. &amp; Brown, K. S. (2004) Phylogeny of the Nymphalidae (Lepidoptera). Systematic Biology, 53, 363 - 383. https: // doi. org / 10.1080 / 10635150490445670" type="journal article" year="2004">Freitas &amp; Brown 2004</bibRefCitation>
: 369), and the justification for its maintenance as a monophyletic group comes from a combined analysis of morphological end molecular data (
<bibRefCitation id="EF924B342865FFF2F44EF8B6FB83B774" author="Wahlberg" box="[809,1056,1841,1866]" pageId="37" pageNumber="38" 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="B977EAD72865FFF2F4C7F8B5FC70B774" box="[928,979,1841,1866]" italics="true" pageId="37" pageNumber="38">et al</emphasis>
. 2009
</bibRefCitation>
). Similarly, even though the nine satyrine tribes recognized in a recent molecular analysis of the subfamily by
<bibRefCitation id="EF924B342865FFF2F488F8D1FAA6B750" author="Pena" box="[1007,1285,1877,1902]" pageId="37" pageNumber="38" refString="Pena, C. &amp; Wahlberg, N. (2008) Prehistorical climate change increased diversification of a group of butterflies. Biology Letters, 4, 274 - 278." type="journal article" year="2008">Peña &amp; Wahlberg (2008)</bibRefCitation>
largely agree with morphological studies, an analysis of the apomorphies of the tribes still is wanting. Thus, opinions of relationship to date have been based on similarity.
</paragraph>
<paragraph id="8BBC36C52865FFF1F7A0F845FB59B32A" blockId="37.[151,1437,833,2010]" lastBlockId="38.[151,1436,151,1365]" lastPageId="38" lastPageNumber="39" pageId="37" pageNumber="38">
On the basis of the wing pattern
<bibRefCitation id="EF924B342865FFF2F550F846FD5CB7E4" author="Nel" box="[567,767,1985,2010]" pageId="37" pageNumber="38" refString="Nel, A. &amp; Nel, J. (1986) A propos de deux Lepidopteres fossiles du Stampien d'Aix- en-Provence (Lepidoptera, Hesperiidae et Nymphalidae Satyrinae), Bulletin du Museum National d' Histoire Naturelle, 8, 343 - 350." type="journal article" year="1986">Nel &amp; Nel (1986)</bibRefCitation>
agree with
<bibRefCitation id="EF924B342865FFF2F4E3F846FBBFB7E4" author="Butler" box="[900,1052,1985,2010]" pageId="37" pageNumber="38" refString="Butler, A. G. (1873) On fossil butterflies. Lepidoptera Exotica, 15, 126 - 128." type="journal article" year="1873">Butler (1873)</bibRefCitation>
and Scudder (1875) about a close relationship with the recent satyrine genus
<taxonomicName id="4C034D462866FFF1F5E4FF1EFD7CB08E" ID-CoL="92GJF" box="[643,735,153,176]" class="Insecta" family="Nymphalidae" genus="Neorina" kingdom="Animalia" order="Lepidoptera" pageId="38" pageNumber="39" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72866FFF1F5E4FF1EFD7CB08E" box="[643,735,153,176]" italics="true" pageId="38" pageNumber="39">Neorina</emphasis>
</taxonomicName>
. They indicate that
<emphasis id="B977EAD72866FFF1F4A2FF1EFBEFB08E" box="[965,1100,153,176]" italics="true" pageId="38" pageNumber="39">Neorinopsis</emphasis>
,
<taxonomicName id="4C034D462866FFF1F33BFF1EFB1BB08E" ID-CoL="92GJF" box="[1116,1208,153,176]" class="Insecta" family="Nymphalidae" genus="Neorina" kingdom="Animalia" order="Lepidoptera" pageId="38" pageNumber="39" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72866FFF1F33BFF1EFB1BB08E" box="[1116,1208,153,176]" italics="true" pageId="38" pageNumber="39">Neorina</emphasis>
</taxonomicName>
and
<taxonomicName id="4C034D462866FFF1F395FF10FF48B0EB" class="Insecta" family="Nymphalidae" genus="Pseudoneorina" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="38" pageNumber="39" phylum="Arthropoda" rank="species" species="coulleti">
<emphasis id="B977EAD72866FFF1F395FF10FF48B0EB" italics="true" pageId="38" pageNumber="39">Pseudoneorina coulleti</emphasis>
</taxonomicName>
(also a fossil species, see under the species name) form a monophyletic group whose ancestor lived in the Eocene or earlier, maybe Upper Cretaceous, f Europe or Asia. As correctly remarked by
<bibRefCitation id="EF924B342866FFF1F3FCFF58FAC8B0C6" author="Nel" box="[1179,1387,223,248]" pageId="38" pageNumber="39" refString="Nel, A. &amp; Nel, J. (1986) A propos de deux Lepidopteres fossiles du Stampien d'Aix- en-Provence (Lepidoptera, Hesperiidae et Nymphalidae Satyrinae), Bulletin du Museum National d' Histoire Naturelle, 8, 343 - 350." type="journal article" year="1986">Nel &amp; Nel (1986)</bibRefCitation>
, the common origin of veins M3 and Cu
<quantity id="4CFB9B202866FFF1F553FE83FDFFB123" box="[564,604,260,285]" metricMagnitude="-2" metricUnit="m" metricValue="2.54" pageId="38" pageNumber="39" unit="in" value="1.0">1 in</quantity>
the forewing is a character found in recent taxa only in the satyrine tribe
<taxonomicName id="4C034D462866FFF1F7F0FEA0FEA4B17E" ID-CoL="93N3Y" box="[151,263,295,320]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="38" pageNumber="39" phylum="Arthropoda" rank="tribe" tribe="Elymniini">Elymniini</taxonomicName>
(Old World Tropics), which is not closely related to
<taxonomicName id="4C034D462866FFF1F403FEAEFC63B17E" ID-CoL="92GJF" box="[868,960,297,320]" class="Insecta" family="Nymphalidae" genus="Neorina" kingdom="Animalia" order="Lepidoptera" pageId="38" pageNumber="39" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72866FFF1F403FEAEFC63B17E" box="[868,960,297,320]" italics="true" pageId="38" pageNumber="39">Neorina</emphasis>
</taxonomicName>
(tribe
<taxonomicName id="4C034D462866FFF1F36AFEA0FBD4B17E" box="[1037,1143,295,320]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="38" pageNumber="39" phylum="Arthropoda" rank="tribe" tribe="Zetherini">Zetherini</taxonomicName>
,
<bibRefCitation id="EF924B342866FFF1F3E4FEAFFA32B17E" author="Pena" box="[1155,1425,295,320]" pageId="38" pageNumber="39" refString="Pena, C. &amp; Wahlberg, N. (2008) Prehistorical climate change increased diversification of a group of butterflies. Biology Letters, 4, 274 - 278." type="journal article" year="2008">Peña &amp; Wahlberg 2008</bibRefCitation>
). Since wing shape in
<taxonomicName id="4C034D462866FFF1F6E5FECBFE51B15B" ID-CoL="93N3Y" box="[386,498,332,357]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="38" pageNumber="39" phylum="Arthropoda" rank="tribe" tribe="Elymniini">Elymniini</taxonomicName>
is generally crenulate and the pattern is very different,
<bibRefCitation id="EF924B342866FFF1F301FECBFA8CB15B" author="Nel" box="[1126,1327,332,357]" pageId="38" pageNumber="39" refString="Nel, A. &amp; Nel, J. (1986) A propos de deux Lepidopteres fossiles du Stampien d'Aix- en-Provence (Lepidoptera, Hesperiidae et Nymphalidae Satyrinae), Bulletin du Museum National d' Histoire Naturelle, 8, 343 - 350." type="journal article" year="1986">Nel &amp; Nel (1986)</bibRefCitation>
conclude that the venational similarity is due to convergence. Alternatively, the similarity of
<taxonomicName id="4C034D462866FFF1F315FEF6FACBB1B6" ID-CoL="92GJF" authority="Neorinopsis" authorityName="Neorinopsis" box="[1138,1384,368,392]" class="Insecta" family="Nymphalidae" genus="Neorina" kingdom="Animalia" order="Lepidoptera" pageId="38" pageNumber="39" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72866FFF1F315FEF6FB6DB1B6" box="[1138,1230,369,392]" italics="true" pageId="38" pageNumber="39">Neorina</emphasis>
,
<emphasis id="B977EAD72866FFF1F387FEF6FACBB1B6" box="[1248,1384,369,392]" italics="true" pageId="38" pageNumber="39">Neorinopsis</emphasis>
</taxonomicName>
and
<taxonomicName id="4C034D462866FFF1F7F0FE13FEE2B193" box="[151,321,404,429]" class="Insecta" family="Nymphalidae" genus="Pseudoneorina" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="38" pageNumber="39" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72866FFF1F7F0FE13FEE2B193" box="[151,321,404,429]" italics="true" pageId="38" pageNumber="39">Pseudoneorina</emphasis>
</taxonomicName>
in wing design could well be based on symplesiomorphy or convergence, and probably the mimetic wing pattern of
<taxonomicName id="4C034D462866FFF1F620FE30FE19B1EE" ID-CoL="93N3Y" box="[327,442,439,464]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="38" pageNumber="39" phylum="Arthropoda" rank="tribe" tribe="Elymniini">Elymniini</taxonomicName>
, mimicking, among others,
<taxonomicName id="4C034D462866FFF1F596FE30FCFDB1EE" box="[753,862,439,464]" class="Insecta" family="Danainae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="38" pageNumber="39" phylum="Arthropoda" rank="family">Danainae</taxonomicName>
, may well have evolved after the common origin of M3 and Cu1 arose in an ancestor of
<emphasis id="B977EAD72866FFF1F522FE5AFD6FB1CA" box="[581,716,477,500]" italics="true" pageId="38" pageNumber="39">Neorinopsis</emphasis>
and
<taxonomicName id="4C034D462866FFF1F46EFE5BFCDEB1CB" ID-CoL="93N3Y" box="[777,893,476,501]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="38" pageNumber="39" phylum="Arthropoda" rank="tribe" tribe="Elymniini">Elymniini</taxonomicName>
. In view of the variability and plasticity in the development of eyespots in
<taxonomicName id="4C034D462866FFF1F6B7FE78FD9EB226" ID-CoL="93MZX" box="[464,573,511,536]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="38" pageNumber="39" phylum="Arthropoda" rank="subFamily" subFamily="Satyrinae">Satyrinae</taxonomicName>
, where even seasonal changes may lead to very different patterns (
<bibRefCitation id="EF924B342866FFF1F243FE78FEB4B203" author="Brakefield" pageId="38" pageNumber="39" refString="Brakefield, P. M., Gates, J., Keys, D., Kesbeke, F., Wijngaarden, P. J., Monteiro, A., French, V. &amp; Carroll, S. B. (1996) Development, plasticity and evolution of butterfly eyespot patterns. Nature, 384, 236 - 242. https: // doi. org / 10.1038 / 384236 a 0" type="journal article" year="1996">
Brakefield
<emphasis id="B977EAD72866FFF1F7F0FDA2FF68B203" box="[151,203,548,573]" italics="true" pageId="38" pageNumber="39">et al</emphasis>
. 1996
</bibRefCitation>
), one should hesitate in providing more weight for similarity in wing design than for similarity in a unique structural character. In this context, the recent publication on Mesozoic kalligrammatid lacewings (Neuroptera) with a surprisingly butterfly-like appearance, including eyespots (
<bibRefCitation id="EF924B342866FFF1F356FDEBFAE3B2BB" author="Labandeira" box="[1073,1344,620,645]" pageId="38" pageNumber="39" refString="Labandeira, C. C., Yang, Q., Santiago-Blay, J. A., Hotton, C. L., Monteiro, A., Wang, Y. - J., Goreva, Y., Shih, C. - K., Siljestrom, S., Rose, T. R., Dilcher, L. D. &amp; Ren, D. (2016) The evolutionary convergence of mid-Mesozoic lacewings and Cenozoic butterflies. Proceedings Royal Society B, 283, 1 - 9." type="journal article" year="2016">
Labandeira
<emphasis id="B977EAD72866FFF1F3DBFDEAFB51B2BB" box="[1212,1266,620,645]" italics="true" pageId="38" pageNumber="39">et al</emphasis>
. 2016
</bibRefCitation>
), which lived several tens of millions of years before the origin of butterflies must be mentioned. This study emphasizes that appearance alone is not a reliable guide to relationship. In an interesting examination, Martins-Neto
<emphasis id="B977EAD72866FFF1F207FD32FA36B2F3" box="[1376,1429,692,717]" italics="true" pageId="38" pageNumber="39">et al</emphasis>
. (1993) described a fossil butterfly from
<collectingCountry id="F31476552866FFF1F533FD50FD34B2CE" box="[596,663,727,752]" name="Brazil" pageId="38" pageNumber="39">Brasil</collectingCountry>
(
<taxonomicName id="4C034D462866FFF1F5C0FD50FCDAB2CE" box="[679,889,727,752]" class="Insecta" family="Nymphalidae" genus="Neorinella" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="38" pageNumber="39" phylum="Arthropoda" rank="species" species="garciae">
<emphasis id="B977EAD72866FFF1F5C0FD50FCDAB2CE" box="[679,889,727,752]" italics="true" pageId="38" pageNumber="39">Neorinella garciae</emphasis>
</taxonomicName>
) with the same remarkable connate origin of M3 and Cu1 from the forewing cell as found in
<emphasis id="B977EAD72866FFF1F5E6FD7AFCAAB32A" box="[641,777,765,788]" italics="true" pageId="38" pageNumber="39">Neorinopsis</emphasis>
and
<taxonomicName id="4C034D462866FFF1F458FD7BFC04B32B" ID-CoL="927WX" box="[831,935,764,789]" class="Insecta" family="Nymphalidae" genus="Elymnias" kingdom="Animalia" order="Lepidoptera" pageId="38" pageNumber="39" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72866FFF1F458FD7BFC04B32B" box="[831,935,764,789]" italics="true" pageId="38" pageNumber="39">Elymnias</emphasis>
</taxonomicName>
(see also
<taxonomicName id="4C034D462866FFF1F371FD7BFB4AB32A" box="[1046,1257,764,789]" class="Insecta" family="Nymphalidae" genus="Neorinella" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="38" pageNumber="39" phylum="Arthropoda" rank="species" species="garciae">
<emphasis id="B977EAD72866FFF1F371FD7BFB4AB32A" box="[1046,1257,764,789]" italics="true" pageId="38" pageNumber="39">Neorinella garciae</emphasis>
</taxonomicName>
).
</paragraph>
<paragraph id="8BBC36C52866FFF1F7A0FC98FC77B49E" blockId="38.[151,1436,151,1365]" pageId="38" pageNumber="39">
<taxonomicName id="4C034D462866FFF1F7A0FC98FED2B306" box="[199,369,799,824]" class="Insecta" family="Nymphalidae" genus="Pseudoneorina" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="38" pageNumber="39" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72866FFF1F7A0FC98FED2B306" box="[199,369,799,824]" italics="true" pageId="38" pageNumber="39">Pseudoneorina</emphasis>
</taxonomicName>
,
<taxonomicName id="4C034D462866FFF1F6E3FC98FE5FB306" box="[388,508,799,824]" class="Insecta" family="Nymphalidae" genus="Neorinella" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="38" pageNumber="39" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72866FFF1F6E3FC98FE5FB306" box="[388,508,799,824]" italics="true" pageId="38" pageNumber="39">Neorinella</emphasis>
</taxonomicName>
,
<taxonomicName id="4C034D462866FFF1F577FCA7FDD1B309" box="[528,626,800,823]" class="Insecta" family="Nymphalidae" genus="Satyrites" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="38" pageNumber="39" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72866FFF1F577FCA7FDD1B309" box="[528,626,800,823]" italics="true" pageId="38" pageNumber="39">Satyrites</emphasis>
</taxonomicName>
and
<emphasis id="B977EAD72866FFF1F5D3FCA6FC98B306" box="[692,827,801,824]" italics="true" pageId="38" pageNumber="39">Neorinopsis</emphasis>
have been classified by
<bibRefCitation id="EF924B342866FFF1F30BFC98FA34B306" author="Wahlberg" box="[1132,1431,799,824]" pageId="38" pageNumber="39" 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>
, without comment, as fossil genera belonging to the subtribe
<taxonomicName id="4C034D462866FFF1F422FCC3FC39B363" box="[837,922,836,861]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="38" pageNumber="39" phylum="Arthropoda" rank="subTribe" subTribe="Lethina">Lethina</taxonomicName>
of the tribe
<taxonomicName id="4C034D462866FFF1F34FFCC3FB20B363" ID-CoL="93N56" box="[1064,1155,836,861]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="38" pageNumber="39" phylum="Arthropoda" rank="tribe" tribe="Satyrini">Satyrini</taxonomicName>
, while
<taxonomicName id="4C034D462866FFF1F3B1FCC2FA91B362" ID-CoL="92GJF" box="[1238,1330,837,860]" class="Insecta" family="Nymphalidae" genus="Neorina" kingdom="Animalia" order="Lepidoptera" pageId="38" pageNumber="39" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72866FFF1F3B1FCC2FA91B362" box="[1238,1330,837,860]" italics="true" pageId="38" pageNumber="39">Neorina</emphasis>
</taxonomicName>
has been placed in the tribe
<taxonomicName id="4C034D462866FFF1F61DFCE0FE47B3BE" box="[378,484,871,896]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="38" pageNumber="39" phylum="Arthropoda" rank="tribe" tribe="Zetherini">Zetherini</taxonomicName>
. For
<emphasis id="B977EAD72866FFF1F542FCEEFD0EB3BE" box="[549,685,873,896]" italics="true" pageId="38" pageNumber="39">Neorinopsis</emphasis>
and
<taxonomicName id="4C034D462866FFF1F58BFCE0FCC7B3BE" box="[748,868,871,896]" class="Insecta" family="Nymphalidae" genus="Neorinella" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="38" pageNumber="39" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72866FFF1F58BFCE0FCC7B3BE" box="[748,868,871,896]" italics="true" pageId="38" pageNumber="39">Neorinella</emphasis>
</taxonomicName>
this placement appears incorrect in view of the venation, and for
<taxonomicName id="4C034D462866FFF1F60EFC0AFE68B39A" box="[361,459,909,932]" class="Insecta" family="Nymphalidae" genus="Satyrites" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="38" pageNumber="39" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72866FFF1F60EFC0AFE68B39A" box="[361,459,909,932]" italics="true" pageId="38" pageNumber="39">Satyrites</emphasis>
</taxonomicName>
it is not certain at all (see the discussion under
<taxonomicName id="4C034D462866FFF1F36BFC0AFBC0B39A" box="[1036,1123,909,932]" class="Insecta" family="Nymphalidae" genus="Satyrites" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="38" pageNumber="39" phylum="Arthropoda" rank="species" species="reynesii">
<emphasis id="B977EAD72866FFF1F36BFC0AFBC0B39A" box="[1036,1123,909,932]" italics="true" pageId="38" pageNumber="39">reynesii</emphasis>
</taxonomicName>
). However, insights in the interrelationships among the tribes of
<taxonomicName id="4C034D462866FFF1F524FC28FD0EB3F6" ID-CoL="93MZX" box="[579,685,943,968]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="38" pageNumber="39" phylum="Arthropoda" rank="subFamily" subFamily="Satyrinae">Satyrinae</taxonomicName>
are still in flux and would be altered with new analyses that apply other or new (molecular) characters and other sets of taxa. Consequently,
<bibRefCitation id="EF924B342866FFF1F4AAFC53FAA2B3D3" author="Wahlberg" box="[973,1281,980,1005]" pageId="38" pageNumber="39" refString="Wahlberg, N. &amp; Brower, A. V. Z. (2006) Satyrinae Boisduval 1833. Version 0 9 November 2006 (under construction). Available from: http: // tolweb. org / Satyrinae / 12189 / 2006.11.09 in The Tree of Life Web Project. Available from: http: // tolweb. org / (accessed 23 May 2015)" type="journal article" year="2006">Wahlberg &amp; Brower (2006)</bibRefCitation>
presented the phylogeny of the tribes of
<taxonomicName id="4C034D462866FFF1F6ADFC70FD97B42E" ID-CoL="93MZX" box="[458,564,1015,1040]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="38" pageNumber="39" phylum="Arthropoda" rank="subFamily" subFamily="Satyrinae">Satyrinae</taxonomicName>
quite differently from the scheme of
<bibRefCitation id="EF924B342866FFF1F481FC7FFAA7B42E" author="Pena" box="[998,1284,1015,1040]" pageId="38" pageNumber="39" refString="Pena, C. &amp; Wahlberg, N. (2008) Prehistorical climate change increased diversification of a group of butterflies. Biology Letters, 4, 274 - 278." type="journal article" year="2008">Peña &amp; Wahlberg (2008)</bibRefCitation>
for the same tribes, and one year later the situation has again changed drastically (
<bibRefCitation id="EF924B342866FFF1F4F3FB9BFBDCB40A" author="Wahlberg" box="[916,1151,1052,1077]" pageId="38" pageNumber="39" 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="B977EAD72866FFF1F360FB9AFB94B40B" box="[1031,1079,1052,1077]" italics="true" pageId="38" pageNumber="39">et al</emphasis>
. 2009
</bibRefCitation>
) (
<figureCitation id="13382A402866FFF1F3F2FB9BFAB4B40A" box="[1173,1303,1052,1077]" captionStart="FIGURES 19 – 21" captionStartId="48.[151,264,605,627]" captionTargetBox="[221,1362,203,576]" captionTargetId="figure@48.[216,1371,193,584]" captionTargetPageId="48" captionText="FIGURES 19 – 21. Three phylogenetic trees for Nymphalidae: Satyrinae. See text for explanation." httpUri="https://zenodo.org/record/583189/files/figure.png" pageId="38" pageNumber="39">Figs 19–21</figureCitation>
). Evidently, if
<emphasis id="B977EAD72866FFF1F7D2FBC6FE38B466" box="[181,411,1087,1112]" italics="true" pageId="38" pageNumber="39">
Neorinopsis
<taxonomicName id="4C034D462866FFF1F62DFBB8FE38B466" box="[330,411,1087,1112]" class="Insecta" family="Nymphalidae" genus="Cyllo" higherTaxonomySource="CoL" kingdom="Animalia" order="Lepidoptera" pageId="38" pageNumber="39" phylum="Arthropoda" rank="species" species="sepulta">sepulta</taxonomicName>
</emphasis>
(about 28–34 Ma) or
<taxonomicName id="4C034D462866FFF1F5C8FBB8FC29B466" box="[687,906,1087,1112]" class="Insecta" family="Nymphalidae" genus="Neorinella" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="38" pageNumber="39" phylum="Arthropoda" rank="species" species="garciae">
<emphasis id="B977EAD72866FFF1F5C8FBB8FC29B466" box="[687,906,1087,1112]" italics="true" pageId="38" pageNumber="39">Neorinella garciae</emphasis>
</taxonomicName>
(about 20–30 Ma) are used to calibrate the molecular clock for
<taxonomicName id="4C034D462866FFF1F619FBE3FE48B443" ID-CoL="93MZX" box="[382,491,1124,1149]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="38" pageNumber="39" phylum="Arthropoda" rank="subFamily" subFamily="Satyrinae">Satyrinae</taxonomicName>
, and the calibration point is placed at the base of
<taxonomicName id="4C034D462866FFF1F341FBE3FB3AB443" ID-CoL="93N3Y" box="[1062,1177,1124,1149]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="38" pageNumber="39" phylum="Arthropoda" rank="tribe" tribe="Elymniini">Elymniini</taxonomicName>
, it makes considerable difference whether it is done in the tree of
<figureCitation id="13382A402866FFF1F514FB00FD61B49E" box="[627,706,1159,1184]" captionStart="FIGURES 19 – 21" captionStartId="48.[151,264,605,627]" captionTargetBox="[221,1362,203,576]" captionTargetId="figure@48.[216,1371,193,584]" captionTargetPageId="48" captionText="FIGURES 19 – 21. Three phylogenetic trees for Nymphalidae: Satyrinae. See text for explanation." httpUri="https://zenodo.org/record/583189/files/figure.png" pageId="38" pageNumber="39">Fig. 19</figureCitation>
or in the tree of
<figureCitation id="13382A402866FFF1F41AFB00FC73B49E" box="[893,976,1159,1184]" captionStart="FIGURES 19 – 21" captionStartId="48.[151,264,605,627]" captionTargetBox="[221,1362,203,576]" captionTargetId="figure@48.[216,1371,193,584]" captionTargetPageId="48" captionText="FIGURES 19 – 21. Three phylogenetic trees for Nymphalidae: Satyrinae. See text for explanation." httpUri="https://zenodo.org/record/583189/files/figure.png" pageId="38" pageNumber="39">Fig. 20</figureCitation>
.
</paragraph>
<paragraph id="8BBC36C52866FFF1F7A0FB2BFEE0B56A" blockId="38.[151,1436,151,1365]" pageId="38" pageNumber="39">
As to geographical relationship, if
<emphasis id="B977EAD72866FFF1F530FB2AFD7DB4FA" box="[599,734,1197,1220]" italics="true" pageId="38" pageNumber="39">Neorinopsis</emphasis>
and
<taxonomicName id="4C034D462866FFF1F47EFB2BFC32B4FB" box="[793,913,1196,1221]" class="Insecta" family="Nymphalidae" genus="Neorinella" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="38" pageNumber="39" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72866FFF1F47EFB2BFC32B4FB" box="[793,913,1196,1221]" italics="true" pageId="38" pageNumber="39">Neorinella</emphasis>
</taxonomicName>
are assigned to
<taxonomicName id="4C034D462866FFF1F329FB2BFB1DB4FB" ID-CoL="93N3Y" box="[1102,1214,1196,1221]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="38" pageNumber="39" phylum="Arthropoda" rank="tribe" tribe="Elymniini">Elymniini</taxonomicName>
on the basis of the connate origin of M3 and Cu1 from the forewing cell,
<taxonomicName id="4C034D462866FFF1F591FB48FCCDB4D6" box="[758,878,1231,1256]" class="Insecta" family="Nymphalidae" genus="Neorinella" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="38" pageNumber="39" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72866FFF1F591FB48FCCDB4D6" box="[758,878,1231,1256]" italics="true" pageId="38" pageNumber="39">Neorinella</emphasis>
</taxonomicName>
would be an example of a fossil found on another continent than the most closely related extant taxa, particularly as
<taxonomicName id="4C034D462866FFF1F410FB73FC44B533" ID-CoL="93N3Y" box="[887,999,1268,1293]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="38" pageNumber="39" phylum="Arthropoda" rank="tribe" tribe="Elymniini">Elymniini</taxonomicName>
currently are restricted to tropical Asia with a single species in Africa (see http://tolweb.org/
<taxonomicName id="4C034D462866FFF1F445FA90FC36B50E" ID-CoL="93N3Y" box="[802,917,1303,1328]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="38" pageNumber="39" phylum="Arthropoda" rank="tribe" tribe="Elymniini">Elymniini</taxonomicName>
/70274). See the section on
<emphasis id="B977EAD72866FFF1F392FA90FE9FB56B" italics="true" pageId="38" pageNumber="39">Biogeographic considerations</emphasis>
.
</paragraph>
</subSubSection>
</treatment>
</document>
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