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<mods:title id="A9137F0ECE48677EC8CC0F86B4456F05">Fossil butterflies, calibration points and the molecular clock (Lepidoptera: Papilionoidea)</mods:title>
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<treatment id="03AA87D3284EFFD8F7F0FB49FB58B0C6" ID-DOI="http://doi.org/10.5281/zenodo.6046907" ID-GBIF-Taxon="130844936" ID-Zenodo-Dep="6046907" LSID="urn:lsid:plazi:treatment:03AA87D3284EFFD8F7F0FB49FB58B0C6" httpUri="http://treatment.plazi.org/id/03AA87D3284EFFD8F7F0FB49FB58B0C6" lastPageId="15" lastPageNumber="15" pageId="14" pageNumber="15">
<subSubSection id="C319654E284EFFD9F7F0FB49FE1AB510" pageId="14" pageNumber="15" type="nomenclature">
<paragraph id="8BBC36C5284EFFD9F7F0FB49FC82B4D6" blockId="14.[151,801,1230,1256]" box="[151,801,1230,1256]" pageId="14" pageNumber="15">
<heading id="D0F481A9284EFFD9F7F0FB49FC82B4D6" bold="true" box="[151,801,1230,1256]" fontSize="11" level="1" pageId="14" pageNumber="15" reason="1">
<emphasis id="B977EAD7284EFFD9F7F0FB49FC82B4D6" bold="true" box="[151,801,1230,1256]" pageId="14" pageNumber="15">
<emphasis id="B977EAD7284EFFD9F7F0FB49FF4EB4D6" bold="true" box="[151,237,1230,1256]" italics="true" pageId="14" pageNumber="15">
<taxonomicName id="4C034D46284EFFD9F7F0FB49FF4BB4D6" box="[151,232,1230,1256]" class="Insecta" family="Nymphalidae" genus="Dynamine" kingdom="Animalia" order="Lepidoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="species" species="alexae">alexae</taxonomicName>
.
</emphasis>
<taxonomicName id="4C034D46284EFFD9F793FB49FC82B4D6" authority="Penalver &amp; Grimaldi, 2006" authorityName="Penalver &amp; Grimaldi" authorityYear="2006" box="[244,801,1230,1256]" class="Insecta" family="Nymphalidae" genus="Dynamine" kingdom="Animalia" order="Lepidoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="species" species="alexae">
<emphasis id="B977EAD7284EFFD9F793FB49FE65B4D6" bold="true" box="[244,454,1230,1256]" italics="true" pageId="14" pageNumber="15">Dynamine alexae</emphasis>
<bibRefCitation id="EF924B34284EFFD9F6A9FB49FC82B4D6" author="Penalver" box="[462,801,1230,1256]" pageId="14" pageNumber="15" refString="Penalver, E. &amp; Grimaldi, D. A. (2006) New data on Miocene butterflies in Dominican Amber (Lerpidoptera: Riodinidae and Nymphalidae) with the description of a new nymphalid. American Museum Novitates, 3591, 1 - 17. https: // doi. org / 10.1206 / 0003 - 0082 (2006) 3519 [1: NDOMBI] 2.0. CO; 2" type="journal article" year="2006">Peñalver &amp; Grimaldi, 2006</bibRefCitation>
</taxonomicName>
</emphasis>
</heading>
</paragraph>
<paragraph id="8BBC36C5284EFFD9F7F0FA92FE1AB510" blockId="14.[151,1437,1301,2010]" box="[151,441,1301,1326]" pageId="14" pageNumber="15">
<taxonomicName id="4C034D46284EFFD9F7F0FA92FE90B510" box="[151,307,1301,1326]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="family">Nymphalidae</taxonomicName>
:
<taxonomicName id="4C034D46284EFFD9F65AFA92FE16B510" box="[317,437,1301,1326]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="subFamily" subFamily="Biblidinae">Biblidinae</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection id="C319654E284EFFD8F7A0FABEFB58B0C6" lastPageId="15" lastPageNumber="16" pageId="14" pageNumber="15" type="description">
<paragraph id="8BBC36C5284EFFD9F7A0FABEFE69B548" blockId="14.[151,1437,1301,2010]" pageId="14" pageNumber="15">
<materialsCitation id="3B6B3C98284EFFD9F7A0FABEFE69B548" ID-GBIF-Occurrence="1503739235" country="Dominican Republic" location="Dominican Amber" municipality="Santago" pageId="14" pageNumber="15" specimenCount="1" stateProvince="Puerto Plata">
<collectingCountry id="F3147655284EFFD9F7A0FABEFE1EB56C" box="[199,445,1337,1362]" name="Dominican Republic" pageId="14" pageNumber="15">Dominican Republic</collectingCountry>
, Cordillera Septentrional between
<collectingMunicipality id="6BD8ACBF284EFFD9F40EFABDFC60B56C" box="[873,963,1338,1362]" pageId="14" pageNumber="15">Santago</collectingMunicipality>
and
<collectingRegion id="49C7F827284EFFD9F368FABDFB07B56C" box="[1039,1188,1337,1362]" country="Dominican Republic" name="Puerto Plata" pageId="14" pageNumber="15">Puerto Plata</collectingRegion>
,
<location id="8EDC601E284EFFD9F3DEFABEFA3BB56C" LSID="urn:lsid:plazi:treatment:03AA87D3284EFFD8F7F0FB49FB58B0C6:8EDC601E284EFFD9F3DEFABEFA3BB56C" box="[1209,1432,1337,1362]" country="Dominican Republic" municipality="Santago" name="Dominican Amber" pageId="14" pageNumber="15" stateProvince="Puerto Plata">Dominican Amber</location>
; Aquitanian, early Miocene.
</materialsCitation>
</paragraph>
<paragraph id="8BBC36C5284EFFD9F7A0FA06FD3FB5A4" blockId="14.[151,1437,1301,2010]" box="[199,668,1409,1434]" pageId="14" pageNumber="15">Depository: AMNH (holotype, DR-18-2).</paragraph>
<paragraph id="8BBC36C5284EFFD9F7A0FA22FCEFB580" blockId="14.[151,1437,1301,2010]" box="[199,844,1445,1470]" pageId="14" pageNumber="15">
Published figures:
<bibRefCitation id="EF924B34284EFFD9F6FEFA22FCEBB580" author="Penalver" box="[409,840,1445,1470]" pageId="14" pageNumber="15" refString="Penalver, E. &amp; Grimaldi, D. A. (2006) New data on Miocene butterflies in Dominican Amber (Lerpidoptera: Riodinidae and Nymphalidae) with the description of a new nymphalid. American Museum Novitates, 3591, 1 - 17. https: // doi. org / 10.1206 / 0003 - 0082 (2006) 3519 [1: NDOMBI] 2.0. CO; 2" type="journal article" year="2006">
Peñalver &amp; Grimaldi (2006:
<figureCitation id="13382A40284EFFD9F5BCFA22FC9DB580" box="[731,830,1445,1470]" captionStart-0="FIGURES 3 4" captionStart-1="FIGURE 5" captionStartId-0="10.[151,264,1801,1823]" captionStartId-1="16.[151,250,1784,1806]" captionTargetBox-0="[165,1429,840,1774]" captionTargetBox-1="[238,1332,193,1743]" captionTargetId-0="figure@10.[154,1438,828,1788]" captionTargetId-1="figure@16.[238,1332,193,1754]" captionTargetPageId-0="10" captionTargetPageId-1="16" captionText-0="FIGURES 3 4. Wing venation of (A) Pareronia Bingham, 1907 (Pieridae), and (B) Hasora Moore, 1881 (Hesperiidae). See text for explanation." captionText-1="FIGURE 5. Ƒanessa amerindica Miller &amp; Brown, 1989, holotype, collections of Invertebrate Paleontology, University of Florida. Photo by Roger Portell. Fig. 6. Pseudoneorina coulleti Nel &amp; Descimon, 1984. France, Dép. Alpes-de-Haute- Provence, Dauphin, specimen in coll. M. Henrotay; photo by courtesy of M. Henrotay. Fig. 7. Archaeolycorea ferreirai Martins-Neto, 1989. Fig. 4 A of original description. Fig. 8. Oligodonta florissantensis Brown, 1976, holotype, collections of Invertebrate Paleontology, University of Florida. Photo by courtesy of Akito Kawahara. Fig. 9. Neorinella garciae Martins- Neto, Kucera-Santos, de Moraes Vieira &amp; de Campos Fragoso, 1993. Fig. 3 A of original description. Fig. 10. Protocoeliades kristenseni de Jong, 2016, holotype, Zoologisk Museum, Copenhagen, Denmark. Photo by Department of Paleobiology, National Museum of Natural History, Washington DC, USA." httpUri-0="https://zenodo.org/record/583186/files/figure.png" httpUri-1="https://zenodo.org/record/583187/files/figure.png" pageId="14" pageNumber="15">Figs 35</figureCitation>
)
</bibRefCitation>
.
</paragraph>
<paragraph id="8BBC36C5284EFFD8F7A0FA4DFB58B0C6" blockId="14.[151,1437,1301,2010]" lastBlockId="15.[151,1436,151,248]" lastPageId="15" lastPageNumber="16" pageId="14" pageNumber="15">
Most of the left wings with the color pattern still partly preserved, abdomen with male genitalia and distal part of mid- and hind legs. The allocation of the fossil to the extant genus
<taxonomicName id="4C034D46284EFFD9F4FFFA69FBA8B63B" box="[920,1035,1518,1541]" class="Insecta" family="Nymphalidae" genus="Dynamine" kingdom="Animalia" order="Lepidoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD7284EFFD9F4FFFA69FBA8B63B" box="[920,1035,1518,1541]" italics="true" pageId="14" pageNumber="15">Dynamine</emphasis>
</taxonomicName>
(
<taxonomicName id="4C034D46284EFFD9F37EFA6AFB17B638" box="[1049,1204,1517,1542]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="family">Nymphalidae</taxonomicName>
,
<taxonomicName id="4C034D46284EFFD9F3D8FA6AFA95B638" box="[1215,1334,1517,1542]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="subFamily" subFamily="Biblidinae">Biblidinae</taxonomicName>
) is based on the striking wing pattern, the dark brown abdomen with a white underside, and the rounded outer margin of the hindwing without expansions or tails. It is similar to the extant genus
<taxonomicName id="4C034D46284EFFD9F4CCF9B1FBA2B673" box="[939,1025,1590,1613]" class="Insecta" family="Nymphalidae" genus="Lucinia" kingdom="Animalia" order="Lepidoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD7284EFFD9F4CCF9B1FBA2B673" box="[939,1025,1590,1613]" italics="true" pageId="14" pageNumber="15">Lucinia</emphasis>
</taxonomicName>
, that however has larger wing ocelli and the margin of the hindwing is not rounded.
<collectingRegion id="49C7F827284EFFD9F5A2F9DDFD44B64F" box="[709,743,1626,1649]" country="Cameroon" name="North" pageId="14" pageNumber="15">No</collectingRegion>
further structural reasons for the allocation of the fossil are given. Unfortunately, the apomorphy of
<taxonomicName id="4C034D46284EFFD9F538F9FAFD77B6A8" box="[607,724,1661,1686]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="subFamily" subFamily="Biblidinae">Biblidinae</taxonomicName>
(the hypandrium) is not mentioned or discussed in
<bibRefCitation id="EF924B34284EFFD9F27BF9FAFEF2B684" author="Penalver" pageId="14" pageNumber="15" refString="Penalver, E. &amp; Grimaldi, D. A. (2006) New data on Miocene butterflies in Dominican Amber (Lerpidoptera: Riodinidae and Nymphalidae) with the description of a new nymphalid. American Museum Novitates, 3591, 1 - 17. https: // doi. org / 10.1206 / 0003 - 0082 (2006) 3519 [1: NDOMBI] 2.0. CO; 2" type="journal article" year="2006">Peñalver &amp; Grimaldi (2006)</bibRefCitation>
, although they extensively discuss the genitalia that are adjacent to the hypandrium (cf.
<bibRefCitation id="EF924B34284EFFD9F254F926FE84B6E0" author="Freitas" pageId="14" pageNumber="15" 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>
). This is regrettable since wing pattern is more likely attributable to convergence and mimicry than to structural characters. This issue is exemplified by the remark by
<bibRefCitation id="EF924B34284EFFD9F408F96EFB0EB73C" author="Penalver" box="[879,1197,1769,1794]" pageId="14" pageNumber="15" refString="Penalver, E. &amp; Grimaldi, D. A. (2006) New data on Miocene butterflies in Dominican Amber (Lerpidoptera: Riodinidae and Nymphalidae) with the description of a new nymphalid. American Museum Novitates, 3591, 1 - 17. https: // doi. org / 10.1206 / 0003 - 0082 (2006) 3519 [1: NDOMBI] 2.0. CO; 2" type="journal article" year="2006">Peñalver &amp; Grimaldi (2006)</bibRefCitation>
that a similar pattern is found in
<taxonomicName id="4C034D46284EFFD9F672F889FEC9B71B" box="[277,362,1806,1829]" class="Insecta" family="Nymphalidae" genus="Lucinia" kingdom="Animalia" order="Lepidoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD7284EFFD9F672F889FEC9B71B" box="[277,362,1806,1829]" italics="true" pageId="14" pageNumber="15">Lucinia</emphasis>
</taxonomicName>
. In the expansive analysis of
<taxonomicName id="4C034D46284EFFD9F5D1F88AFCEEB718" box="[694,845,1805,1830]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="family">Nymphalidae</taxonomicName>
by
<bibRefCitation id="EF924B34284EFFD9F411F88AFBD7B718" box="[886,1140,1805,1830]" pageId="14" pageNumber="15" 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="B977EAD7284EFFD9F48EF889FBBAB718" box="[1001,1049,1805,1830]" italics="true" pageId="14" pageNumber="15">et al</emphasis>
. (2009)
</bibRefCitation>
, the subfamily is split into two monophyletic groups. In
<bibRefCitation id="EF924B34284EFFD9F686F8B6FD7DB774" box="[481,734,1841,1866]" pageId="14" pageNumber="15" 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="B977EAD7284EFFD9F533F8B5FD27B774" box="[596,644,1841,1866]" italics="true" pageId="14" pageNumber="15">et al</emphasis>
. (2009)
</bibRefCitation>
<taxonomicName id="4C034D46284EFFD9F583F8B5FCF4B777" box="[740,855,1842,1865]" class="Insecta" family="Nymphalidae" genus="Dynamine" kingdom="Animalia" order="Lepidoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD7284EFFD9F583F8B5FCF4B777" box="[740,855,1842,1865]" italics="true" pageId="14" pageNumber="15">Dynamine</emphasis>
</taxonomicName>
and
<taxonomicName id="4C034D46284EFFD9F4E9F8B5FC40B777" box="[910,995,1842,1865]" class="Insecta" family="Nymphalidae" genus="Lucinia" kingdom="Animalia" order="Lepidoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD7284EFFD9F4E9F8B5FC40B777" box="[910,995,1842,1865]" italics="true" pageId="14" pageNumber="15">Lucinia</emphasis>
</taxonomicName>
are in the same group, but distant from each other, with
<taxonomicName id="4C034D46284EFFD9F631F8D1FE6AB753" box="[342,457,1878,1901]" class="Insecta" family="Nymphalidae" genus="Dynamine" kingdom="Animalia" order="Lepidoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD7284EFFD9F631F8D1FE6AB753" box="[342,457,1878,1901]" italics="true" pageId="14" pageNumber="15">Dynamine</emphasis>
</taxonomicName>
being the first offshoot of the group (of which 18 genera have been included), while
<taxonomicName id="4C034D46284EFFD9F7F0F8FDFF4EB7AF" box="[151,237,1914,1937]" class="Insecta" family="Nymphalidae" genus="Lucinia" kingdom="Animalia" order="Lepidoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD7284EFFD9F7F0F8FDFF4EB7AF" box="[151,237,1914,1937]" italics="true" pageId="14" pageNumber="15">Lucinia</emphasis>
</taxonomicName>
is five divergence events later nearer to the crown. These relationships suggest that their similarity is either due to convergence, or alternatively to a symplesiomorphy. In the absence of structural arguments, and if the fossil is used for calibration, then it should be placed near the root of the
<taxonomicName id="4C034D46284EFFD9F4E0F846FBA3B7E4" box="[903,1024,1985,2010]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="subFamily" subFamily="Biblidinae">Biblidinae</taxonomicName>
. This is precisely where
<bibRefCitation id="EF924B34284EFFD8F272F846FEAAB08E" lastPageId="15" lastPageNumber="16" pageId="14" pageNumber="15" 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="B977EAD7284EFFD8F2EFF845FF0EB08E" italics="true" lastPageId="15" lastPageNumber="16" pageId="14" pageNumber="15">et al</emphasis>
. (2009)
</bibRefCitation>
put the calibration point. However, they put a minimum date on this point of 20 Ma. According to
<bibRefCitation id="EF924B34284FFFD8F7F0FF3BFD92B0EB" author="Iturralde-Vinent" box="[151,561,188,213]" pageId="15" pageNumber="16" refString="Iturralde-Vinent, M. A. &amp; MacPhee, R. D. E. (1996) Age and paleographical origin of Dominican amber. Science, 273, 1850 - 1852." type="journal article" year="1996">Iturralde-Vinent &amp; MacPhee (1996)</bibRefCitation>
, the age of Dominican amber is 1520 Ma. By choosing the higher age, the derived divergence estimate may be 33 % too high, if 15 Ma would be the correct age for the amber.
</paragraph>
</subSubSection>
</treatment>
</document>