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<document id="F1DCCCBFA4BEADCF12603AB4612CF70C" 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="03AA87D32855FFC0F7F0F897FE13B1CB" 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="Lethe" docType="treatment" docVersion="9" lastPageNumber="23" masterDocId="FF93FFAB2840FFD7F767FF87FFA3B03E" masterDocTitle="Fossil butterflies, calibration points and the molecular clock (Lepidoptera: Papilionoidea)" masterLastPageNumber="63" masterPageNumber="1" pageNumber="22" updateTime="1698439901205" updateUser="ExternalLinkService">
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<mods:title id="DDBAA41588212AE49ED744EBE601B464">Fossil butterflies, calibration points and the molecular clock (Lepidoptera: Papilionoidea)</mods:title>
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<mods:namePart id="53D055C1724CB58AC32322B37A41EE0A">Jong, Rienk De</mods:namePart>
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<mods:date id="4EF779F5A1AF5C80C027AD25018120B1">2017</mods:date>
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<mods:identifier id="CBD471D071828758C862C78C1323B64F" type="CLB-Dataset">32975</mods:identifier>
<mods:identifier id="DE081CB3FA573AD336D2757C16324C58" type="DOI">10.5281/zenodo.583183</mods:identifier>
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<treatment id="03AA87D32855FFC0F7F0F897FE13B1CB" ID-DOI="http://doi.org/10.5281/zenodo.6046938" ID-GBIF-Taxon="130844924" ID-Zenodo-Dep="6046938" LSID="urn:lsid:plazi:treatment:03AA87D32855FFC0F7F0F897FE13B1CB" httpUri="http://treatment.plazi.org/id/03AA87D32855FFC0F7F0F897FE13B1CB" lastPageId="23" lastPageNumber="23" pageId="21" pageNumber="22">
<subSubSection id="C319654E2855FFC2F7F0F897FCAAB714" box="[151,777,1808,1834]" pageId="21" pageNumber="22" type="nomenclature">
<paragraph id="8BBC36C52855FFC2F7F0F897FCAAB714" blockId="21.[151,777,1808,1834]" box="[151,777,1808,1834]" pageId="21" pageNumber="22">
<heading id="D0F481A92855FFC2F7F0F897FCAAB714" bold="true" box="[151,777,1808,1834]" fontSize="11" level="1" pageId="21" pageNumber="22" reason="1">
<emphasis id="B977EAD72855FFC2F7F0F897FCAAB714" bold="true" box="[151,777,1808,1834]" pageId="21" pageNumber="22">
<emphasis id="B977EAD72855FFC2F7F0F897FE74B714" bold="true" box="[151,471,1808,1834]" italics="true" pageId="21" pageNumber="22">
corbieri.
<taxonomicName id="4C034D462855FFC2F663F897FEEAB714" ID-CoL="92D6C" box="[260,329,1808,1834]" class="Insecta" family="Nymphalidae" genus="Lethe" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="genus">Lethe</taxonomicName>
(?) corbieri
</emphasis>
<bibRefCitation id="EF924B342855FFC2F6B8F897FCA7B714" author="Nel" box="[479,772,1808,1834]" pageId="21" pageNumber="22" refString="Nel, A., Nel, J. &amp; Balme, C. (1993) Un nouveau Lepidoptere Satyrinae fossile de l'Oligocene du Sud-Est de la France (Insecta, Lepidoptera, Nymphalidae). Linneana belgica, 14, 20 - 36." type="journal article" year="1993">Nel, Nel &amp; Balme, 1993</bibRefCitation>
.
</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="C319654E2855FFC0F7F0F8D1FE13B1CB" lastPageId="23" lastPageNumber="24" pageId="21" pageNumber="22" type="description">
<paragraph id="8BBC36C52855FFC2F7F0F8D1FC04B751" blockId="21.[151,1221,1878,2011]" box="[151,935,1878,1903]" pageId="21" pageNumber="22">
<taxonomicName id="4C034D462855FFC2F7F0F8D1FE90B751" box="[151,307,1878,1903]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="family">Nymphalidae</taxonomicName>
:
<taxonomicName id="4C034D462855FFC2F65AF8D1FE04B751" box="[317,423,1878,1903]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="subFamily" subFamily="Satyrinae">Satyrinae</taxonomicName>
(subdivisions not yet generally agreed upon).
</paragraph>
<paragraph id="8BBC36C52855FFC2F7A0F8FCFC21B7AD" blockId="21.[151,1221,1878,2011]" box="[199,898,1914,1939]" pageId="21" pageNumber="22">
<materialsCitation id="3B6B3C982855FFC2F7A0F8FCFC21B7AD" ID-GBIF-Occurrence="1503739214" box="[199,898,1914,1939]" collectorName="Dep. de Vaucluse" country="France" location="France" pageId="21" pageNumber="22" specimenCount="1">
<collectingCountry id="F31476552855FFC2F7A0F8FCFEB5B7AC" box="[199,278,1915,1938]" name="France" pageId="21" pageNumber="22">France</collectingCountry>
,
<collectorName id="26F653132855FFC2F646F8FCFE44B7AD" box="[289,487,1914,1939]" pageId="21" pageNumber="22">Dép. de Vaucluse</collectorName>
, Céreste; Rupelian, early Oligocene.
</materialsCitation>
</paragraph>
<paragraph id="8BBC36C52855FFC2F7A0F819FB09B7E5" blockId="21.[151,1221,1878,2011]" pageId="21" pageNumber="22">
<materialsCitation id="3B6B3C982855FFC2F7A0F819FB09B7E5" ID-GBIF-Occurrence="1503739223" collectorName="Musee du Parc Naturel Regional du Luberon" country="France" location="France" pageId="21" pageNumber="22" specimenCount="1" typeStatus="holotype">
Depository:
<typeStatus id="54B888672855FFC2F635F819FE14B789" box="[338,439,1950,1975]" pageId="21" pageNumber="22" type="holotype">holotype</typeStatus>
,
<collectorName id="26F653132855FFC2F6A5F818FC18B789" box="[450,955,1950,1975]" pageId="21" pageNumber="22">Musée du Parc Naturel Régional du Lubéron</collectorName>
, Apt, Vaucluse,
<collectingCountry id="F31476552855FFC2F316F818FB62B788" box="[1137,1217,1951,1974]" name="France" pageId="21" pageNumber="22">France</collectingCountry>
. Published figures:
<bibRefCitation id="EF924B342855FFC2F6FEF845FD12B7E5" author="Nel" box="[409,689,1986,2011]" pageId="21" pageNumber="22" refString="Nel, A., Nel, J. &amp; Balme, C. (1993) Un nouveau Lepidoptere Satyrinae fossile de l'Oligocene du Sud-Est de la France (Insecta, Lepidoptera, Nymphalidae). Linneana belgica, 14, 20 - 36." type="journal article" year="1993">Nel, Nel &amp; Balme (1993</bibRefCitation>
:
<figureCitation id="13382A402855FFC2F5DBF845FC86B7E5" box="[700,805,1986,2011]" captionStart-0="FIGURE 1" captionStart-1="FIGURE 2" captionStart-2="FIGURES 3 – 4" captionStartId-0="4.[151,250,1051,1073]" captionStartId-1="7.[151,250,1650,1672]" captionStartId-2="10.[151,264,1801,1823]" captionTargetBox-0="[167,1414,420,1017]" captionTargetBox-1="[175,1415,420,1615]" captionTargetBox-2="[165,1429,840,1774]" captionTargetId-0="figure@4.[157,1429,411,1026]" captionTargetId-1="figure@7.[151,1436,409,1627]" captionTargetId-2="figure@10.[154,1438,828,1788]" captionTargetPageId-0="4" captionTargetPageId-1="7" captionTargetPageId-2="10" captionText-0="FIGURE 1. Localities of fossil butterflies plotted ion a modern map. See Table 1 for further, complementary information." captionText-1="FIGURE 2. Apomorphies, their placement on a tree, and the assignment of calibration points. See text for explanation." captionText-2="FIGURES 3 – 4. Wing venation of (A) Pareronia Bingham, 1907 (Pieridae), and (B) Hasora Moore, 1881 (Hesperiidae). See text for explanation." httpUri-0="https://zenodo.org/record/583184/files/figure.png" httpUri-1="https://zenodo.org/record/583185/files/figure.png" httpUri-2="https://zenodo.org/record/583186/files/figure.png" pageId="21" pageNumber="22">Figs. 1–3</figureCitation>
);?
<bibRefCitation id="EF924B342855FFC2F428F845FB05B7E5" author="Pfretzschner" box="[847,1190,1986,2011]" pageId="21" pageNumber="22" refString="Pfretzschner, H. U. (1998) Ein weitertes Exemplar von Lethe? corbieri Nel 1993 (Lepidoptera, Satyridae) aus dem Unter- Oligozan von Cereste (Sud-Frankreich). Palaontologische Zeitschrift, 72, 59 - 64." type="journal article" year="1998">
Pfretzschner (1998:
<figureCitation id="13382A402855FFC2F355F845FB3FB7E5" box="[1074,1180,1986,2011]" captionStart-0="FIGURE 1" captionStart-1="FIGURE 2" captionStart-2="FIGURES 3 – 4" captionStartId-0="4.[151,250,1051,1073]" captionStartId-1="7.[151,250,1650,1672]" captionStartId-2="10.[151,264,1801,1823]" captionTargetBox-0="[167,1414,420,1017]" captionTargetBox-1="[175,1415,420,1615]" captionTargetBox-2="[165,1429,840,1774]" captionTargetId-0="figure@4.[157,1429,411,1026]" captionTargetId-1="figure@7.[151,1436,409,1627]" captionTargetId-2="figure@10.[154,1438,828,1788]" captionTargetPageId-0="4" captionTargetPageId-1="7" captionTargetPageId-2="10" captionText-0="FIGURE 1. Localities of fossil butterflies plotted ion a modern map. See Table 1 for further, complementary information." captionText-1="FIGURE 2. Apomorphies, their placement on a tree, and the assignment of calibration points. See text for explanation." captionText-2="FIGURES 3 – 4. Wing venation of (A) Pareronia Bingham, 1907 (Pieridae), and (B) Hasora Moore, 1881 (Hesperiidae). See text for explanation." httpUri-0="https://zenodo.org/record/583184/files/figure.png" httpUri-1="https://zenodo.org/record/583185/files/figure.png" httpUri-2="https://zenodo.org/record/583186/files/figure.png" pageId="21" pageNumber="22">Figs. 1–3</figureCitation>
)
</bibRefCitation>
.
</materialsCitation>
</paragraph>
<paragraph id="8BBC36C52856FFC1F7A0FF1FFE07B1CB" blockId="22.[151,1437,151,2013]" pageId="22" pageNumber="23">
A nicely preserved specimen, probably female. Antennae tricarinate (autapomorphy of
<taxonomicName id="4C034D462856FFC1F3B2FF10FACCB08E" box="[1237,1391,151,176]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="22" pageNumber="23" phylum="Arthropoda" rank="family">Nymphalidae</taxonomicName>
, cf.
<bibRefCitation id="EF924B342856FFC1F7F0FF3BFED1B0EA" author="Ackery" box="[151,370,188,213]" pageId="22" pageNumber="23" 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>
), club weakly developed. Eyes hairy. Forelegs reduced (as in all
<taxonomicName id="4C034D462856FFC1F300FF3BFB5DB0EB" box="[1127,1278,188,213]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="22" pageNumber="23" phylum="Arthropoda" rank="family">Nymphalidae</taxonomicName>
and males of most
<taxonomicName id="4C034D462856FFC1F7B1FF58FEF8B0C6" box="[214,347,223,248]" class="Insecta" family="Lycaenidae" kingdom="Animalia" order="Lepidoptera" pageId="22" pageNumber="23" phylum="Arthropoda" rank="family">Lycaenidae</taxonomicName>
). Forewing with vein 2A missing; radial formula 1, 2, 3+(4+5); R-branches and M1 originating very closely together from upper corner of cell; cell closed, about half as long as costa; Sc and 2A are weakly but distinctly swollen at their base; outer margin indented between M3 and Cu1 (or apically produced); prominent white-centred eye-spot between M1 and M2, and a small dark dot directly beneath it. Hindwing venation without special characters; cell closed; veins M3 and Cu1 originating from the same point (as in many
<taxonomicName id="4C034D462856FFC1F3D3FEE8FAF3B1B6" box="[1204,1360,367,392]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="22" pageNumber="23" phylum="Arthropoda" rank="family">Nymphalidae</taxonomicName>
); outer margin indented between M2 and M3; in the original description it is interpreted as a slight extension at the end of vein M3 reminiscent of a rudimentary tail. Rather well preserved pattern of eye-spots and a submarginal line on forewing and hindwing.
</paragraph>
<paragraph id="8BBC36C52856FFC1F7A0FD87FEABB296" blockId="22.[151,1437,151,2013]" pageId="22" pageNumber="23">
A swollen base of Sc and 2A in the forewing is an apomorphy of
<taxonomicName id="4C034D462856FFC1F4D6FE78FBBDB226" box="[945,1054,511,536]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="22" pageNumber="23" phylum="Arthropoda" rank="subFamily" subFamily="Satyrinae">Satyrinae</taxonomicName>
. It is, however, not universal, and absent in genera like
<taxonomicName id="4C034D462856FFC1F6EEFDA3FE50B203" box="[393,499,548,573]" class="Insecta" family="Nymphalidae" genus="Melanitis" kingdom="Animalia" order="Lepidoptera" pageId="22" pageNumber="23" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72856FFC1F6EEFDA3FE50B203" box="[393,499,548,573]" italics="true" pageId="22" pageNumber="23">Melanitis</emphasis>
</taxonomicName>
and
<taxonomicName id="4C034D462856FFC1F54CFDA3FDC9B203" box="[555,618,548,573]" class="Insecta" family="Nymphalidae" genus="Lethe" kingdom="Animalia" order="Lepidoptera" pageId="22" pageNumber="23" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72856FFC1F54CFDA3FDC9B203" box="[555,618,548,573]" italics="true" pageId="22" pageNumber="23">Lethe</emphasis>
</taxonomicName>
(
<bibRefCitation id="EF924B342856FFC1F51EFDA3FCF2B202" author="Ackery" box="[633,849,548,573]" pageId="22" pageNumber="23" 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="B977EAD72856FFC1F5B2FDA2FCA5B203" box="[725,774,548,573]" italics="true" pageId="22" pageNumber="23">et al</emphasis>
. 1999
</bibRefCitation>
; see also below). Moreover, a swollen base of Sc is also found in various other nymphalid genera, as mentioned below, under
<taxonomicName id="4C034D462856FFC1F48AFDCFFB13B25E" box="[1005,1200,584,608]" class="Insecta" family="Nymphalidae" genus="Satyrites" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="22" pageNumber="23" phylum="Arthropoda" rank="species" species="reynesii">
<emphasis id="B977EAD72856FFC1F48AFDCFFB13B25E" box="[1005,1200,584,608]" italics="true" pageId="22" pageNumber="23">Satyrites reynesii</emphasis>
</taxonomicName>
, but in that case the forewing cell is open, or practically so. The eyespot between M1 and M2 also is widespread but not universal in
<taxonomicName id="4C034D462856FFC1F7F0FD08FEA6B296" box="[151,261,655,680]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="22" pageNumber="23" phylum="Arthropoda" rank="subFamily" subFamily="Satyrinae">Satyrinae</taxonomicName>
.
</paragraph>
<paragraph id="8BBC36C52856FFC1F7A0FD33FC30B49E" blockId="22.[151,1437,151,2013]" pageId="22" pageNumber="23">
The authors discussed the fossil at length and compared it with the information in the monograph of
<bibRefCitation id="EF924B342856FFC1F231FD33FF46B2CE" author="Miller" pageId="22" pageNumber="23" 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>
. They arrived at the conclusion that it most probably belongs to the “groupe de
<taxonomicName id="4C034D462856FFC1F33EFD50FB34B2CE" box="[1113,1175,727,752]" class="Insecta" family="Nymphalidae" genus="Lethe" kingdom="Animalia" order="Lepidoptera" pageId="22" pageNumber="23" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72856FFC1F33EFD50FB34B2CE" box="[1113,1175,727,752]" italics="true" pageId="22" pageNumber="23">Lethe</emphasis>
</taxonomicName>
”, at present in the tribe
<taxonomicName id="4C034D462856FFC1F7F0FD7BFF50B32B" box="[151,243,764,789]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="22" pageNumber="23" phylum="Arthropoda" rank="tribe" tribe="Satyrini">Satyrini</taxonomicName>
. Since they could not identify the fossil more precisely, &quot;nous nous contenterons de le placer dans le genre
<emphasis id="B977EAD72856FFC1F7F0FC98FEEFB306" box="[151,332,799,824]" italics="true" pageId="22" pageNumber="23">
<taxonomicName id="4C034D462856FFC1F7F0FC98FF76B306" box="[151,213,799,824]" class="Insecta" family="Nymphalidae" genus="Lethe" kingdom="Animalia" order="Lepidoptera" pageId="22" pageNumber="23" phylum="Arthropoda" rank="genus" sensu="lato">Lethe</taxonomicName>
<taxonomicNameLabel id="A24457AC2856FFC1F7BBFCA6FEEFB306" box="[220,332,799,824]" pageId="22" pageNumber="23" sensu="lato">sensu lato</taxonomicNameLabel>
</emphasis>
&quot; (&quot;we must be satisfied with placing it in the genus …&quot;), as an indication that the fossil belongs to the group of genera considered closely related to
<taxonomicName id="4C034D462856FFC1F5DDFCC3FD5BB363" box="[698,760,836,861]" class="Insecta" family="Nymphalidae" genus="Lethe" kingdom="Animalia" order="Lepidoptera" pageId="22" pageNumber="23" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72856FFC1F5DDFCC3FD5BB363" box="[698,760,836,861]" italics="true" pageId="22" pageNumber="23">Lethe</emphasis>
</taxonomicName>
by
<bibRefCitation id="EF924B342856FFC1F446FCC3FC1FB363" author="Miller" box="[801,956,836,861]" pageId="22" pageNumber="23" 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>
. In their nomenclature they tried to express this by naming the fossil &quot;
<emphasis id="B977EAD72856FFC1F6A4FCE0FD60B3BE" box="[451,707,871,896]" italics="true" pageId="22" pageNumber="23">
<taxonomicName id="4C034D462856FFC1F6A4FCE0FDA2B3BE" box="[451,513,871,896]" class="Insecta" family="Nymphalidae" genus="Lethe" kingdom="Animalia" order="Lepidoptera" pageId="22" pageNumber="23" phylum="Arthropoda" rank="genus">Lethe</taxonomicName>
(?) corbieri n.sp
</emphasis>
.&quot; However, the &quot;groupe de
<taxonomicName id="4C034D462856FFC1F365FCE0FBE3B3BE" box="[1026,1088,871,896]" class="Insecta" family="Nymphalidae" genus="Lethe" kingdom="Animalia" order="Lepidoptera" pageId="22" pageNumber="23" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72856FFC1F365FCE0FBE3B3BE" box="[1026,1088,871,896]" italics="true" pageId="22" pageNumber="23">Lethe</emphasis>
</taxonomicName>
&quot;, apparently the same as what was called &quot;
<taxonomicName id="4C034D462856FFC1F644FC0BFEC2B39B" box="[291,353,908,933]" class="Insecta" family="Nymphalidae" genus="Lethe" kingdom="Animalia" order="Lepidoptera" pageId="22" pageNumber="23" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72856FFC1F644FC0BFEC2B39B" box="[291,353,908,933]" italics="true" pageId="22" pageNumber="23">Lethe</emphasis>
</taxonomicName>
-series,
<emphasis id="B977EAD72856FFC1F6DDFC0AFDEAB39A" box="[442,585,909,932]" italics="true" pageId="22" pageNumber="23">sensu stricto</emphasis>
&quot; by
<bibRefCitation id="EF924B342856FFC1F5E4FC0BFCBDB39B" author="Miller" box="[643,798,908,933]" pageId="22" pageNumber="23" 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>
, contains 26 nominal genera that nowadays are divided among at least two tribes (
<taxonomicName id="4C034D462856FFC1F6A3FC28FDBCB3F6" box="[452,543,943,968]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="22" pageNumber="23" phylum="Arthropoda" rank="tribe" tribe="Satyrini">Satyrini</taxonomicName>
and
<taxonomicName id="4C034D462856FFC1F53FFC28FD69B3F6" box="[600,714,943,968]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="22" pageNumber="23" phylum="Arthropoda" rank="tribe" tribe="Elymniini">Elymniini</taxonomicName>
) in the classification followed by the Natural History Museum, London, in their catalogue of butterflies and moths of the world (http://www.nhm.ac.uk/our-science/data/butmoth/), which is based on the spreadsheet of Lamas_Genera_
<date id="FFBD10052856FFC1F461FC70FCFFB42E" box="[774,860,1015,1040]" pageId="22" pageNumber="23" value="08-04-11">041108</date>
.xls. Moreover, the use of the genus name, even with a question mark, suggests a closer relationship than may be real. In extant
<taxonomicName id="4C034D462856FFC1F4BCFB9BFBB9B40B" box="[987,1050,1052,1077]" class="Insecta" family="Nymphalidae" genus="Lethe" kingdom="Animalia" order="Lepidoptera" pageId="22" pageNumber="23" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72856FFC1F4BCFB9BFBB9B40B" box="[987,1050,1052,1077]" italics="true" pageId="22" pageNumber="23">Lethe</emphasis>
</taxonomicName>
there are no basally swollen veins in the forewing (
<bibRefCitation id="EF924B342856FFC1F634FBB8FD88B466" author="Ackery" box="[339,555,1087,1112]" pageId="22" pageNumber="23" 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="B977EAD72856FFC1F6C8FBC6FE43B466" box="[431,480,1087,1112]" italics="true" pageId="22" pageNumber="23">et al</emphasis>
. 1999
</bibRefCitation>
;
<bibRefCitation id="EF924B342856FFC1F550FBB8FC93B466" author="Schatz" box="[567,816,1087,1112]" pageId="22" pageNumber="23" refString="Schatz, E. &amp; Rober, J. (1892) Die Familien und Gattungen der Tagfalter Systematisch und Analytisch Bearbeitet. Lowensohn, Furth, ii + 284 pp." type="book" year="1892">Schatz &amp; Röber 1892</bibRefCitation>
: Pl. 33); at most, Sc is thicker in its proximal part than the radius, or fused with the proximal part of the radius to a thicker vein (see, e.g.,
<bibRefCitation id="EF924B342856FFC1F34DFBE3FABDB442" author="Bascombe" box="[1066,1310,1124,1149]" pageId="22" pageNumber="23" refString="Bascombe, M. J., Johnston, G. &amp; Bascombe, F. S. (1999) The Butterflies of Hong Kong. Academic Press, London, ix + 422 pp." type="book" year="1999">
Bascombe
<emphasis id="B977EAD72856FFC1F3C2FBE2FB77B443" box="[1189,1236,1124,1149]" italics="true" pageId="22" pageNumber="23">et al</emphasis>
. 1999
</bibRefCitation>
, Fig. 9.10). It is uncertain if such a subtle difference is clearly visible in a fossil.
</paragraph>
<paragraph id="8BBC36C52856FFC1F7A0FB2AFB29B773" blockId="22.[151,1437,151,2013]" pageId="22" pageNumber="23">
<bibRefCitation id="EF924B342856FFC1F7A0FB2AFE4EB4FA" author="Pena" box="[199,493,1196,1221]" pageId="22" pageNumber="23" 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>
used the fossil for calibrating the molecular clock of their phylogenetic tree of
<taxonomicName id="4C034D462856FFC1F7F0FB48FEA7B4D6" box="[151,260,1231,1256]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="22" pageNumber="23" phylum="Arthropoda" rank="subFamily" subFamily="Satyrinae">Satyrinae</taxonomicName>
, placing it at the point of divergence of
<taxonomicName id="4C034D462856FFC1F5A0FB48FCA5B4D6" box="[711,774,1231,1256]" class="Insecta" family="Nymphalidae" genus="Lethe" kingdom="Animalia" order="Lepidoptera" pageId="22" pageNumber="23" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72856FFC1F5A0FB48FCA5B4D6" box="[711,774,1231,1256]" italics="true" pageId="22" pageNumber="23">Lethe</emphasis>
</taxonomicName>
and
<taxonomicName id="4C034D462856FFC1F45AFB56FC26B4D6" box="[829,901,1233,1256]" class="Insecta" family="Nymphalidae" genus="Neope" kingdom="Animalia" order="Lepidoptera" pageId="22" pageNumber="23" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72856FFC1F45AFB56FC26B4D6" box="[829,901,1233,1256]" italics="true" pageId="22" pageNumber="23">Neope</emphasis>
</taxonomicName>
, the only two genera of Miller's (1968) &quot;
<taxonomicName id="4C034D462856FFC1F233FB48FA31B4D6" box="[1364,1426,1231,1256]" class="Insecta" family="Nymphalidae" genus="Lethe" kingdom="Animalia" order="Lepidoptera" pageId="22" pageNumber="23" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72856FFC1F233FB48FA31B4D6" box="[1364,1426,1231,1256]" italics="true" pageId="22" pageNumber="23">Lethe</emphasis>
</taxonomicName>
- series,
<emphasis id="B977EAD72856FFC1F782FB72FED0B532" box="[229,371,1269,1292]" italics="true" pageId="22" pageNumber="23">sensu stricto</emphasis>
&quot; included in their analysis.
<bibRefCitation id="EF924B342856FFC1F5D7FB73FC0CB532" box="[688,943,1268,1293]" pageId="22" pageNumber="23" 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">
Wahlberg
<emphasis id="B977EAD72856FFC1F443FB72FCFEB533" box="[804,861,1268,1293]" italics="true" pageId="22" pageNumber="23">et al.</emphasis>
(2009)
</bibRefCitation>
included more genera of this group in their analysis of the
<taxonomicName id="4C034D462856FFC1F627FA90FE78B50E" box="[320,475,1303,1328]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="22" pageNumber="23" phylum="Arthropoda" rank="family">Nymphalidae</taxonomicName>
.
<taxonomicName id="4C034D462856FFC1F682FA90FD80B50E" box="[485,547,1303,1328]" class="Insecta" family="Nymphalidae" genus="Lethe" kingdom="Animalia" order="Lepidoptera" pageId="22" pageNumber="23" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72856FFC1F682FA90FD80B50E" box="[485,547,1303,1328]" italics="true" pageId="22" pageNumber="23">Lethe</emphasis>
</taxonomicName>
ended up as sister to the genera
<taxonomicName id="4C034D462856FFC1F4E8FA90FC5FB50E" box="[911,1020,1303,1328]" class="Insecta" family="Nymphalidae" genus="Satyrodes" higherTaxonomySource="CoL" kingdom="Animalia" order="Lepidoptera" pageId="22" pageNumber="23" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72856FFC1F4E8FA90FC5FB50E" box="[911,1020,1303,1328]" italics="true" pageId="22" pageNumber="23">Satyrodes</emphasis>
</taxonomicName>
and
<taxonomicName id="4C034D462856FFC1F353FA90FB26B50E" box="[1076,1157,1303,1328]" class="Insecta" family="Nymphalidae" genus="Enodia" kingdom="Animalia" order="Lepidoptera" pageId="22" pageNumber="23" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72856FFC1F353FA90FB26B50E" box="[1076,1157,1303,1328]" italics="true" pageId="22" pageNumber="23">Enodia</emphasis>
</taxonomicName>
combined; together they were sister to the genus
<taxonomicName id="4C034D462856FFC1F6C4FABBFD85B56B" box="[419,550,1340,1365]" class="Insecta" family="Nymphalidae" genus="Rhaphicera" kingdom="Animalia" order="Lepidoptera" pageId="22" pageNumber="23" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72856FFC1F6C4FABBFD85B56B" box="[419,550,1340,1365]" italics="true" pageId="22" pageNumber="23">Rhaphicera</emphasis>
</taxonomicName>
, which in Miller's (1968) study was closer to the &quot;
<taxonomicName id="4C034D462856FFC1F330FABAFB10B56A" box="[1111,1203,1341,1364]" class="Insecta" family="Nymphalidae" genus="Pararge" kingdom="Animalia" order="Lepidoptera" pageId="22" pageNumber="23" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72856FFC1F330FABAFB10B56A" box="[1111,1203,1341,1364]" italics="true" pageId="22" pageNumber="23">Pararge</emphasis>
</taxonomicName>
-series&quot;, and the four genera together were sister to
<taxonomicName id="4C034D462856FFC1F68AFAE6FD97B546" box="[493,564,1377,1400]" class="Insecta" family="Nymphalidae" genus="Neope" kingdom="Animalia" order="Lepidoptera" pageId="22" pageNumber="23" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72856FFC1F68AFAE6FD97B546" box="[493,564,1377,1400]" italics="true" pageId="22" pageNumber="23">Neope</emphasis>
</taxonomicName>
. In this study the fossil was used to constrain the split between
<taxonomicName id="4C034D462856FFC1F26CFAD8FAE9B546" box="[1291,1354,1375,1400]" class="Insecta" family="Nymphalidae" genus="Lethe" kingdom="Animalia" order="Lepidoptera" pageId="22" pageNumber="23" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72856FFC1F26CFAD8FAE9B546" box="[1291,1354,1375,1400]" italics="true" pageId="22" pageNumber="23">Lethe</emphasis>
</taxonomicName>
and its sister group (
<taxonomicName id="4C034D462856FFC1F649FA03FE38B5A3" box="[302,411,1412,1437]" class="Insecta" family="Nymphalidae" genus="Satyrodes" higherTaxonomySource="CoL" kingdom="Animalia" order="Lepidoptera" pageId="22" pageNumber="23" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72856FFC1F649FA03FE38B5A3" box="[302,411,1412,1437]" italics="true" pageId="22" pageNumber="23">Satyrodes</emphasis>
</taxonomicName>
+
<taxonomicName id="4C034D462856FFC1F6CCFA03FE5FB5A3" box="[427,508,1412,1437]" class="Insecta" family="Nymphalidae" genus="Enodia" kingdom="Animalia" order="Lepidoptera" pageId="22" pageNumber="23" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72856FFC1F6CCFA03FE5FB5A3" box="[427,508,1412,1437]" italics="true" pageId="22" pageNumber="23">Enodia</emphasis>
</taxonomicName>
) to the age of the fossil (supposed to be 25 Ma). This is not the same calibration point as in
<bibRefCitation id="EF924B342856FFC1F671FA2FFD97B5FE" author="Pena" box="[278,564,1447,1472]" pageId="22" pageNumber="23" 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>
. Supposing the relationship as found in
<bibRefCitation id="EF924B342856FFC1F49CFA20FB59B5FE" box="[1019,1274,1447,1472]" pageId="22" pageNumber="23" 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">
Wahlberg
<emphasis id="B977EAD72856FFC1F309FA2EFB02B5FE" box="[1134,1185,1447,1472]" italics="true" pageId="22" pageNumber="23">et al</emphasis>
. (2009)
</bibRefCitation>
is correct, the calibration point in
<bibRefCitation id="EF924B342856FFC1F610FA4AFD30B5DA" author="Pena" box="[375,659,1484,1509]" pageId="22" pageNumber="23" 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>
is too low down in the tree. Since the number of substitutions since the split between
<taxonomicName id="4C034D462856FFC1F63DFA76FE01B636" box="[346,418,1521,1544]" class="Insecta" family="Nymphalidae" genus="Neope" kingdom="Animalia" order="Lepidoptera" pageId="22" pageNumber="23" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72856FFC1F63DFA76FE01B636" box="[346,418,1521,1544]" italics="true" pageId="22" pageNumber="23">Neope</emphasis>
</taxonomicName>
and its sister group must have been greater than since the split of
<taxonomicName id="4C034D462856FFC1F31AFA68FB18B636" box="[1149,1211,1519,1544]" class="Insecta" family="Nymphalidae" genus="Lethe" kingdom="Animalia" order="Lepidoptera" pageId="22" pageNumber="23" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72856FFC1F31AFA68FB18B636" box="[1149,1211,1519,1544]" italics="true" pageId="22" pageNumber="23">Lethe</emphasis>
</taxonomicName>
and its sister group, the substitution rate was higher, consisting of more substitutions over the same period), favoring the calibration point of
<bibRefCitation id="EF924B342856FFC1F667F9BFFD84B66E" author="Pena" box="[256,551,1591,1616]" pageId="22" pageNumber="23" 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>
rather than with the calibration point of
<bibRefCitation id="EF924B342856FFC1F371F9B0FA87B66E" box="[1046,1316,1591,1616]" pageId="22" pageNumber="23" 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">
Wahlberg
<emphasis id="B977EAD72856FFC1F3EAF9BEFB60B66E" box="[1165,1219,1591,1616]" italics="true" pageId="22" pageNumber="23">et al</emphasis>
. (2009)
</bibRefCitation>
. A higher substitution rate leads to lower divergence time estimates. If all 10 genera of the subtribe
<taxonomicName id="4C034D462856FFC1F3CEF9DBFB5DB64B" box="[1193,1278,1628,1653]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="22" pageNumber="23" phylum="Arthropoda" rank="subTribe" subTribe="Lethina">Lethina</taxonomicName>
(
<bibRefCitation id="EF924B342856FFC1F268F9DBFEADB6A6" author="Wahlberg" pageId="22" pageNumber="23" 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>
) had been included in the analysis, the outcome may have again been different. Whether in this case the difference would be significant, remains to be seen. I just draw attention to a) the importance of finding the correct position of the fossil on the tree, and b) the large impact of the tree itself, for making reliable estimates of divergence times. Since there is still so much uncertainty about the phylogeny of the subfamily and the exact position of the fossil on the tree is difficult to ascertain, this fossil can, for the time being, only be used as minimum age calibration point at the root of the
<taxonomicName id="4C034D462856FFC1F523F8B3FD12B773" box="[580,689,1844,1869]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="22" pageNumber="23" phylum="Arthropoda" rank="subFamily" subFamily="Satyrinae">Satyrinae</taxonomicName>
, based on similarities rather apomorphies.
</paragraph>
<paragraph id="8BBC36C52856FFC0F7A0F8D0FE13B1CB" blockId="22.[151,1437,151,2013]" lastBlockId="23.[151,1437,151,501]" lastPageId="23" lastPageNumber="24" pageId="22" pageNumber="23">
<bibRefCitation id="EF924B342856FFC1F7A0F8D0FE0BB74E" author="Pfretzschner" box="[199,424,1879,1904]" pageId="22" pageNumber="23" refString="Pfretzschner, H. U. (1998) Ein weitertes Exemplar von Lethe? corbieri Nel 1993 (Lepidoptera, Satyridae) aus dem Unter- Oligozan von Cereste (Sud-Frankreich). Palaontologische Zeitschrift, 72, 59 - 64." type="journal article" year="1998">Pfretzschner (1998)</bibRefCitation>
recorded a partly damaged forewing from Céreste as belonging to the same taxon. He again described (and figured) the basally swollen Sc and 2A, and the eyespot between M1 and M
<quantity id="4CFB9B202856FFC1F3B1F8FBFAA3B7AB" box="[1238,1280,1916,1941]" metricMagnitude="-2" metricUnit="m" metricValue="5.08" pageId="22" pageNumber="23" unit="in" value="2.0">2 in</quantity>
the forewing. Since the length of the cell (as far as can be deduced from what is visible of the veins, and extrapolating the front margin and termen to the probable position of the apex) is
<emphasis id="B977EAD72856FFC1F449F842FCF3B7E2" box="[814,848,1989,2012]" italics="true" pageId="22" pageNumber="23">ca.</emphasis>
40 % of the length of the wing), whereas it is about half the length of the wing in
<emphasis id="B977EAD72857FFC0F696FF10FD1EB08E" box="[497,701,151,176]" italics="true" pageId="23" pageNumber="24">
<taxonomicName id="4C034D462857FFC0F696FF10FD8CB08E" box="[497,559,151,176]" class="Insecta" family="Nymphalidae" genus="Lethe" kingdom="Animalia" order="Lepidoptera" pageId="23" pageNumber="24" phylum="Arthropoda" rank="genus">Lethe</taxonomicName>
(?) corbieri
</emphasis>
, it could well be the remnant of a different taxon. According to
<bibRefCitation id="EF924B342857FFC0F7F0FF3BFED5B0EA" author="Pfretzschner" box="[151,374,188,213]" pageId="23" pageNumber="24" refString="Pfretzschner, H. U. (1998) Ein weitertes Exemplar von Lethe? corbieri Nel 1993 (Lepidoptera, Satyridae) aus dem Unter- Oligozan von Cereste (Sud-Frankreich). Palaontologische Zeitschrift, 72, 59 - 64." type="journal article" year="1998">Pfretzschner (1998)</bibRefCitation>
the wing design and the uniform pigmentation of the wing surface are identical to
<emphasis id="B977EAD72857FFC0F253FF3BFF52B0C6" italics="true" pageId="23" pageNumber="24">
<taxonomicName id="4C034D462857FFC0F253FF3BFAD1B0EB" box="[1332,1394,188,213]" class="Insecta" family="Nymphalidae" genus="Lethe" kingdom="Animalia" order="Lepidoptera" pageId="23" pageNumber="24" phylum="Arthropoda" rank="genus">Lethe</taxonomicName>
(?) corbieri
</emphasis>
; but since the fossil is so fragmentary, and the external similarity of many extant satyrine taxa can be confusing, identification of this fossil with
<emphasis id="B977EAD72857FFC0F5F7FE83FCFDB123" box="[656,862,260,285]" italics="true" pageId="23" pageNumber="24">
<taxonomicName id="4C034D462857FFC0F5F7FE83FD6DB123" box="[656,718,260,285]" class="Insecta" family="Nymphalidae" genus="Lethe" kingdom="Animalia" order="Lepidoptera" pageId="23" pageNumber="24" phylum="Arthropoda" rank="genus">Lethe</taxonomicName>
(?) corbieri
</emphasis>
seems premature. See also the discussion on the plasticity of wing pattern under
<emphasis id="B977EAD72857FFC0F57EFEAEFD03B17E" box="[537,672,297,320]" italics="true" pageId="23" pageNumber="24">Neorinopsis</emphasis>
<taxonomicName id="4C034D462857FFC0F5CAFEA0FC31B17E" authority="Boisduval" authorityName="Boisduval" box="[685,914,295,320]" class="Insecta" family="Nymphalidae" genus="Cyllo" higherTaxonomySource="CoL" kingdom="Animalia" order="Lepidoptera" pageId="23" pageNumber="24" phylum="Arthropoda" rank="species" species="sepulta">
<emphasis id="B977EAD72857FFC0F5CAFEA0FD5DB17E" box="[685,766,295,320]" italics="true" pageId="23" pageNumber="24">sepulta</emphasis>
(Boisduval)
</taxonomicName>
. Moreover, in the satyrine tribes
<taxonomicName id="4C034D462857FFC0F244FEA0FA34B17E" box="[1315,1431,295,320]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="23" pageNumber="24" phylum="Arthropoda" rank="tribe" tribe="Brassolini">Brassolini</taxonomicName>
,
<taxonomicName id="4C034D462857FFC0F7F0FECBFEA2B15B" box="[151,257,332,357]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="23" pageNumber="24" phylum="Arthropoda" rank="tribe" tribe="Morphini">Morphini</taxonomicName>
and
<taxonomicName id="4C034D462857FFC0F65CFECBFE6BB15B" box="[315,456,332,357]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="23" pageNumber="24" phylum="Arthropoda" rank="tribe" tribe="Amathusiini">Amathusiini</taxonomicName>
, the eyespot of the forewing may be present on the underside only, and since the thin chitinous wing surface may be completely lost in the fossil and only the melanin may be preserved (see
<bibRefCitation id="EF924B342857FFC0F7F0FE13FEC5B192" author="Pfretzschner" box="[151,358,404,429]" pageId="23" pageNumber="24" refString="Pfretzschner, H. U. (1998) Ein weitertes Exemplar von Lethe? corbieri Nel 1993 (Lepidoptera, Satyridae) aus dem Unter- Oligozan von Cereste (Sud-Frankreich). Palaontologische Zeitschrift, 72, 59 - 64." type="journal article" year="1998">Pfretzschner 1998</bibRefCitation>
for chemical background; see also
<bibRefCitation id="EF924B342857FFC0F469FE13FBBBB193" author="Labandeira" box="[782,1048,404,429]" pageId="23" pageNumber="24" 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="B977EAD72857FFC0F4F0FE12FC6FB193" box="[919,972,404,429]" italics="true" pageId="23" pageNumber="24">et al</emphasis>
. 2016
</bibRefCitation>
for detection of melanin-bearing eyespots in much older insects), it cannot be decided whether the eyespot in the living insect was on the upperside or underside of the wing.
</paragraph>
</subSubSection>
</treatment>
</document>
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