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<mods:title id="17187C2F95C94B3B7819017DE635612B">Fossil butterflies, calibration points and the molecular clock (Lepidoptera: Papilionoidea)</mods:title>
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<treatment id="03AA87D32858FFCEF7F0F8D7FBBBB2F3" ID-DOI="http://doi.org/10.5281/zenodo.6046948" ID-GBIF-Taxon="130844865" ID-Zenodo-Dep="6046948" LSID="urn:lsid:plazi:treatment:03AA87D32858FFCEF7F0F8D7FBBBB2F3" httpUri="http://treatment.plazi.org/id/03AA87D32858FFCEF7F0F8D7FBBBB2F3" lastPageId="25" lastPageNumber="26" pageId="24" pageNumber="25">
<subSubSection id="C319654E2858FFCFF7F0F8D7FD0FB792" pageId="24" pageNumber="25" type="nomenclature">
<paragraph id="8BBC36C52858FFCFF7F0F8D7FCE6B754" blockId="24.[151,837,1872,1898]" box="[151,837,1872,1898]" pageId="24" pageNumber="25">
<heading id="D0F481A92858FFCFF7F0F8D7FCE6B754" bold="true" box="[151,837,1872,1898]" fontSize="11" level="1" pageId="24" pageNumber="25" reason="1">
<emphasis id="B977EAD72858FFCFF7F0F8D7FCE6B754" bold="true" box="[151,837,1872,1898]" pageId="24" pageNumber="25">
<emphasis id="B977EAD72858FFCFF7F0F8D7FF58B754" bold="true" box="[151,251,1872,1898]" italics="true" pageId="24" pageNumber="25">
<taxonomicName id="4C034D462858FFCFF7F0F8D7FF54B754" box="[151,247,1872,1898]" class="Insecta" family="Riodinidae" genus="Voltinia" kingdom="Animalia" order="Lepidoptera" pageId="24" pageNumber="25" phylum="Arthropoda" rank="species" species="dramba">dramba</taxonomicName>
.
</emphasis>
<taxonomicName id="4C034D462858FFCFF665F8D7FCE6B754" authority="Hall, Robbins &amp; Harvey, 2004" authorityName="Hall, Robbins &amp; Harvey" authorityYear="2004" box="[258,837,1872,1898]" class="Insecta" family="Riodinidae" genus="Voltinia" kingdom="Animalia" order="Lepidoptera" pageId="24" pageNumber="25" phylum="Arthropoda" rank="species" species="dramba">
<emphasis id="B977EAD72858FFCFF665F8D7FE66B754" bold="true" box="[258,453,1872,1898]" italics="true" pageId="24" pageNumber="25">Voltinia dramba</emphasis>
<bibRefCitation id="EF924B342858FFCFF6ABF8D7FCE6B754" author="Hall" box="[460,837,1872,1898]" pageId="24" pageNumber="25" refString="Hall, J. P. W., Robbins, R. K. &amp; Harvey, D. J. (2004) Extinction and biogeography in the Caribbean: new evidence from a fossil riodinid butterfly in Dominican amber. Proceedings of the Royal Society of London B, 271, 797 - 801. https: // doi. org / 10.1098 / rspb. 2004.2691" type="journal article" year="2004">Hall, Robbins &amp; Harvey, 2004</bibRefCitation>
</taxonomicName>
</emphasis>
</heading>
</paragraph>
<paragraph id="8BBC36C52858FFCFF7F0F813FD0FB792" blockId="24.[151,1436,1940,2037]" box="[151,684,1940,1965]" pageId="24" pageNumber="25">
<taxonomicName id="4C034D462858FFCFF7F0F813FEB4B793" box="[151,279,1940,1965]" class="Insecta" family="Riodinidae" kingdom="Animalia" order="Lepidoptera" pageId="24" pageNumber="25" phylum="Arthropoda" rank="family">Riodinidae</taxonomicName>
:
<taxonomicName id="4C034D462858FFCFF647F813FE03B793" box="[288,416,1940,1965]" class="Insecta" family="Riodininae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="24" pageNumber="25" phylum="Arthropoda" rank="family">Riodininae</taxonomicName>
:
<taxonomicName id="4C034D462858FFCFF6CDF813FD9CB793" box="[426,575,1940,1965]" class="Insecta" family="Riodinidae" kingdom="Animalia" order="Lepidoptera" pageId="24" pageNumber="25" phylum="Arthropoda" rank="tribe" tribe="Mesosemiini">Mesosemiini</taxonomicName>
:
<emphasis id="B977EAD72858FFCFF52DF813FD05B793" box="[586,678,1940,1965]" italics="true" pageId="24" pageNumber="25">Voltinea</emphasis>
.
</paragraph>
</subSubSection>
<subSubSection id="C319654E2858FFCEF7A0F83FFBBBB2F3" lastPageId="25" lastPageNumber="26" pageId="24" pageNumber="25" type="description">
<paragraph id="8BBC36C52858FFCFF7A0F83FFE71B7CB" blockId="24.[151,1436,1940,2037]" pageId="24" pageNumber="25">
<materialsCitation id="3B6B3C982858FFCFF7A0F83FFE71B7CB" ID-GBIF-Occurrence="1503739229" country="Dominican Republic" location="Dominican" pageId="24" pageNumber="25" specimenCount="1" stateProvince="Santiago">
<collectingCountry id="F31476552858FFCFF7A0F83FFE1EB7EF" box="[199,445,1976,2001]" name="Dominican Republic" pageId="24" pageNumber="25">Dominican Republic</collectingCountry>
, Cordillera Septentrional between
<collectingRegion id="49C7F8272858FFCFF40EF83FFC68B7EF" box="[873,971,1976,2001]" country="Dominican Republic" name="Santiago" pageId="24" pageNumber="25">Santiago</collectingRegion>
and
<collectingRegion id="49C7F8272858FFCFF371F83EFB0FB7EF" box="[1046,1196,1976,2001]" country="Dominican Republic" name="Puerto Plata" pageId="24" pageNumber="25">Puerto Plata</collectingRegion>
,
<location id="8EDC601E2858FFCFF3A6F83FFA9EB7EF" LSID="urn:lsid:plazi:treatment:03AA87D32858FFCEF7F0F8D7FBBBB2F3:8EDC601E2858FFCFF3A6F83FFA9EB7EF" box="[1217,1341,1976,2001]" country="Dominican Republic" name="Dominican" pageId="24" pageNumber="25" stateProvince="Santiago">Dominican</location>
amber; Burdigalian, early Miocene.
</materialsCitation>
</paragraph>
<paragraph id="8BBC36C52859FFCEF7A0FF10FD39B0EA" blockId="25.[151,1436,151,717]" pageId="25" pageNumber="26">
Depositories: USNM (holotype);
<materialsCitation id="3B6B3C982859FFCEF538FF1FFB91B08E" ID-GBIF-Occurrence="1503739234" box="[607,1074,151,176]" collectorName="Murata" country="Japan" location="Kyota" pageId="25" pageNumber="26" specimenCount="1" typeStatus="paratype">
<collectorName id="26F653132859FFCEF538FF1FFD13B091" box="[607,688,152,175]" pageId="25" pageNumber="26">Murata</collectorName>
coll.,
<location id="8EDC601E2859FFCEF46BFF1FFCF0B08E" LSID="urn:lsid:plazi:treatment:03AA87D32858FFCEF7F0F8D7FBBBB2F3:8EDC601E2859FFCEF46BFF1FFCF0B08E" box="[780,851,152,176]" country="Japan" name="Kyota" pageId="25" pageNumber="26">Kyota</location>
,
<collectingCountry id="F31476552859FFCEF40EFF1FFC0AB091" box="[873,937,152,175]" name="Japan" pageId="25" pageNumber="26">Japan</collectingCountry>
(
<typeStatus id="54B888672859FFCEF4A4FF1FFB89B08E" box="[963,1066,152,176]" pageId="25" pageNumber="26" type="paratype">paratype</typeStatus>
)
</materialsCitation>
;
<materialsCitation id="3B6B3C982859FFCEF32FFF1FFD39B0EA" ID-GBIF-Occurrence="1503739213" collectionCode="AMNH" collectorName="Morrone" country="Italy" location="Torino" pageId="25" pageNumber="26" specimenCount="1" typeStatus="paratype">
<collectorName id="26F653132859FFCEF32FFF1FFB09B091" box="[1096,1194,152,175]" pageId="25" pageNumber="26">Morrone</collectorName>
coll.,
<location id="8EDC601E2859FFCEF261FF10FAF0B08E" LSID="urn:lsid:plazi:treatment:03AA87D32858FFCEF7F0F8D7FBBBB2F3:8EDC601E2859FFCEF261FF10FAF0B08E" box="[1286,1363,151,176]" country="Italy" name="Torino" pageId="25" pageNumber="26">Torino</location>
,
<collectingCountry id="F31476552859FFCEF20EFF10FA3FB08E" box="[1385,1436,151,176]" name="Italy" pageId="25" pageNumber="26">Italy</collectingCountry>
(
<typeStatus id="54B888672859FFCEF7F8FF3BFEA5B0EA" box="[159,262,188,212]" pageId="25" pageNumber="26" type="paratype">paratype</typeStatus>
);
<collectionCode id="ED12AE002859FFCEF67EFF3BFECDB0EA" box="[281,366,188,212]" country="USA" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34925" name="American Museum of Natural History" pageId="25" pageNumber="26">AMNH</collectionCode>
(one specimen, DR-18-1).
</materialsCitation>
</paragraph>
<paragraph id="8BBC36C52859FFCEF7A0FF58FBCFB0C6" blockId="25.[151,1436,151,717]" box="[199,1132,223,248]" pageId="25" pageNumber="26">
Published figures:
<bibRefCitation id="EF924B342859FFCEF6FEFF58FDF7B0C6" author="Hall" box="[409,596,223,248]" pageId="25" pageNumber="26" refString="Hall, J. P. W., Robbins, R. K. &amp; Harvey, D. J. (2004) Extinction and biogeography in the Caribbean: new evidence from a fossil riodinid butterfly in Dominican amber. Proceedings of the Royal Society of London B, 271, 797 - 801. https: // doi. org / 10.1098 / rspb. 2004.2691" type="journal article" year="2004">
Hall
<emphasis id="B977EAD72859FFCEF6B7FF66FDA2B0C6" box="[464,513,223,248]" italics="true" pageId="25" pageNumber="26">et al</emphasis>
. (2004
</bibRefCitation>
:
<figureCitation id="13382A402859FFCEF538FF58FD3CB0C6" box="[607,671,223,248]" captionStart="FIGURE 1" captionStartId="4.[151,250,1051,1073]" captionTargetBox="[167,1414,420,1017]" captionTargetId="figure@4.[157,1429,411,1026]" captionTargetPageId="4" captionText="FIGURE 1. Localities of fossil butterflies plotted ion a modern map. See Table 1 for further, complementary information." httpUri="https://zenodo.org/record/583184/files/figure.png" pageId="25" pageNumber="26">Fig. 1</figureCitation>
);
<bibRefCitation id="EF924B342859FFCEF5DEFF58FBCAB0C6" author="Penalver" box="[697,1129,223,248]" pageId="25" pageNumber="26" 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="13382A402859FFCEF49CFF58FBE3B0C6" box="[1019,1088,223,248]" captionStart="FIGURE 1" captionStartId="4.[151,250,1051,1073]" captionTargetBox="[167,1414,420,1017]" captionTargetId="figure@4.[157,1429,411,1026]" captionTargetPageId="4" captionText="FIGURE 1. Localities of fossil butterflies plotted ion a modern map. See Table 1 for further, complementary information." httpUri="https://zenodo.org/record/583184/files/figure.png" pageId="25" pageNumber="26">Figs 1</figureCitation>
,
<figureCitation id="13382A402859FFCEF329FF67FBFCB0C6" box="[1102,1119,224,248]" 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="25" pageNumber="26">2</figureCitation>
)
</bibRefCitation>
.
</paragraph>
<paragraph id="8BBC36C52859FFCEF7A0FE83FBBBB2F3" blockId="25.[151,1436,151,717]" pageId="25" pageNumber="26">
The fossil was described by
<bibRefCitation id="EF924B342859FFCEF560FE83FD6AB122" box="[519,713,260,285]" pageId="25" pageNumber="26" refString="Hall, J. P. W., Robbins, R. K. &amp; Harvey, D. J. (2004) Extinction and biogeography in the Caribbean: new evidence from a fossil riodinid butterfly in Dominican amber. Proceedings of the Royal Society of London B, 271, 797 - 801. https: // doi. org / 10.1098 / rspb. 2004.2691" type="journal article">
Hall
<emphasis id="B977EAD72859FFCEF559FE82FDCCB123" box="[574,623,260,285]" italics="true" pageId="25" pageNumber="26">et al</emphasis>
. (2004)
</bibRefCitation>
as a member of the extant genus
<taxonomicName id="4C034D462859FFCEF325FE83FA8CB123" box="[1090,1327,260,285]" class="Insecta" family="Riodinidae" genus="Voltinia" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72859FFCEF325FE83FB3AB123" box="[1090,1177,260,285]" italics="true" pageId="25" pageNumber="26">Voltinia</emphasis>
(Riodinidae)
</taxonomicName>
after five female specimens &quot;that collectively represent probably the best-preserved fossil record for any lepidopteran.&quot; An additional fossil specimen belonging to the same taxon was described by
<bibRefCitation id="EF924B342859FFCEF4B4FECBFAB6B15B" author="Penalver" box="[979,1301,332,357]" pageId="25" pageNumber="26" 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>
. Because of the exceptionally good quality of the fossils, there is little reason to doubt its taxonomic position, the more so since
<bibRefCitation id="EF924B342859FFCEF7F0FE13FEBBB192" author="Hall" box="[151,280,404,429]" pageId="25" pageNumber="26" refString="Hall, J. P. W. (2003) Phylogenetic reassessment of the five forewing radial-veined tribes of Riodininae (Lepidoptera: Riodinidae). Systematic Entomology, 28, 23 - 37." type="journal article" year="2003">Hall (2003)</bibRefCitation>
had published a phylogenetic reassessment based on morphology of the tribes of
<taxonomicName id="4C034D462859FFCEF3CEFE13FA84B193" box="[1193,1319,404,429]" class="Insecta" family="Riodininae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="family">Riodininae</taxonomicName>
, the genus
<taxonomicName id="4C034D462859FFCEF7F0FE30FF4DB1EE" box="[151,238,439,464]" class="Insecta" family="Riodinidae" genus="Voltinia" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72859FFCEF7F0FE30FF4DB1EE" box="[151,238,439,464]" italics="true" pageId="25" pageNumber="26">Voltinia</emphasis>
</taxonomicName>
belongs to, and thus, characters of all related extant genera were well known and evaluated. Very recently an extensive molecular phylogenetic analysis of the family was published (
<bibRefCitation id="EF924B342859FFCEF36DFE5BFAA3B1CB" author="Espeland" box="[1034,1280,476,501]" pageId="25" pageNumber="26" refString="Espeland, M., Hall, J. P. W., DeVries, P. J., Lees, D. C., Cornwall, M., Hsu, Y-F., Wu, L-W., Campbell, D. L., Talavera, G., Vila, R., Salzman, S., Ruehr, S., Lohman, D. J. &amp; Pierce, N. (2015) Ancient Neotropical origin and recent recolonization: phylogeny, biogeography and diversification of the Riodinidae (Lepidoptera: Papilionoidea). Molecular Phylogenetics and Evolution, 93, 296 - 306." type="journal article" year="2015">
Espeland
<emphasis id="B977EAD72859FFCEF319FE5AFB10B1CB" box="[1150,1203,476,501]" italics="true" pageId="25" pageNumber="26">et al</emphasis>
. 2015
</bibRefCitation>
) covering all subfamilies, tribes and subtribes, and 98 out of 145 described genera of the
<taxonomicName id="4C034D462859FFCEF36EFE78FB24B226" box="[1033,1159,511,536]" class="Insecta" family="Riodinidae" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="family">Riodinidae</taxonomicName>
, including
<taxonomicName id="4C034D462859FFCEF26FFE78FAFCB226" box="[1288,1375,511,536]" class="Insecta" family="Riodinidae" genus="Voltinia" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72859FFCEF26FFE78FAFCB226" box="[1288,1375,511,536]" italics="true" pageId="25" pageNumber="26">Voltinia</emphasis>
</taxonomicName>
. The taxonomic position of the fossil was used as one of the calibration points (placed at the common stem of
<taxonomicName id="4C034D462859FFCEF222FDA3FA3FB203" box="[1349,1436,548,573]" class="Insecta" family="Riodinidae" genus="Voltinia" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72859FFCEF222FDA3FA3FB203" box="[1349,1436,548,573]" italics="true" pageId="25" pageNumber="26">Voltinia</emphasis>
</taxonomicName>
and
<taxonomicName id="4C034D462859FFCEF7A1FDCEFEBEB25E" box="[198,285,585,608]" class="Insecta" family="Lycaenidae" genus="Napaea" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72859FFCEF7A1FDCEFEBEB25E" box="[198,285,585,608]" italics="true" pageId="25" pageNumber="26">Napaea</emphasis>
</taxonomicName>
), with an age of 1525 Ma as given by
<bibRefCitation id="EF924B342859FFCEF5B8FDC0FC07B25E" box="[735,932,583,608]" pageId="25" pageNumber="26" refString="Hall, J. P. W., Robbins, R. K. &amp; Harvey, D. J. (2004) Extinction and biogeography in the Caribbean: new evidence from a fossil riodinid butterfly in Dominican amber. Proceedings of the Royal Society of London B, 271, 797 - 801. https: // doi. org / 10.1098 / rspb. 2004.2691" type="journal article">
Hall
<emphasis id="B977EAD72859FFCEF471FDCEFCEBB25E" box="[790,840,583,608]" italics="true" pageId="25" pageNumber="26">et al</emphasis>
. (2004)
</bibRefCitation>
. Since the Burdigalian lasted from 20.43 Ma to 15.97 Ma, the latter (younger) age should be used for calibration. However, recent studies, soon to be published, suggest that the allocation of the fossil needs revision (Noemy Seraphim Pereira, pers. comm.), and it may be wise to postpone the use of the fossil for calibration until its position is better known.
</paragraph>
</subSubSection>
</treatment>
</document>