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<mods:title id="0C3D35E186A50A2C5B7DABC2BB3F27BC">Fossil butterflies, calibration points and the molecular clock (Lepidoptera: Papilionoidea)</mods:title>
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<treatment id="03AA87D32868FFFFF7F0FF1FFADBB444" ID-DOI="http://doi.org/10.5281/zenodo.6047074" ID-GBIF-Taxon="130844858" ID-Zenodo-Dep="6047074" LSID="urn:lsid:plazi:treatment:03AA87D32868FFFFF7F0FF1FFADBB444" httpUri="http://treatment.plazi.org/id/03AA87D32868FFFFF7F0FF1FFADBB444" lastPageNumber="41" pageId="40" pageNumber="41">
<subSubSection id="C319654E2868FFFFF7F0FF1FFC42B0C9" pageId="40" pageNumber="41" type="nomenclature">
<paragraph id="8BBC36C52868FFFFF7F0FF1FFD59B08C" blockId="40.[151,762,152,178]" box="[151,762,152,178]" pageId="40" pageNumber="41">
<heading id="D0F481A92868FFFFF7F0FF1FFD59B08C" bold="true" box="[151,762,152,178]" fontSize="11" level="1" pageId="40" pageNumber="41" reason="1">
<emphasis id="B977EAD72868FFFFF7F0FF1FFD59B08C" bold="true" box="[151,762,152,178]" pageId="40" pageNumber="41">
<emphasis id="B977EAD72868FFFFF7F0FF1FFE87B08C" bold="true" box="[151,292,152,178]" italics="true" pageId="40" pageNumber="41">
<taxonomicName id="4C034D462868FFFFF7F0FF1FFEBCB08C" box="[151,287,152,178]" class="Insecta" family="Edelfalter" genus="Prolibythea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="40" pageNumber="41" phylum="Arthropoda" rank="species" species="vagabunda">vagabunda</taxonomicName>
.
</emphasis>
<taxonomicName id="4C034D462868FFFFF64CFF1FFD59B08C" authority="Scudder, 1889" authorityName="Scudder" authorityYear="1889" box="[299,762,152,178]" class="Insecta" family="Edelfalter" genus="Prolibythea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="40" pageNumber="41" phylum="Arthropoda" rank="species" species="vagabunda">
<emphasis id="B977EAD72868FFFFF64CFF1FFDE7B08C" bold="true" box="[299,580,152,178]" italics="true" pageId="40" pageNumber="41">Prolibythea vagabunda</emphasis>
<bibRefCitation id="EF924B342868FFFFF52CFF1FFD59B08C" author="Scudder" box="[587,762,152,178]" pageId="40" pageNumber="41" refString="Scudder, S. H. (1889) The Fossil butterflies of Florissant. United States Geological Survey, 8 th Annual Report, 439 - 472." type="book chapter" year="1889">Scudder, 1889</bibRefCitation>
</taxonomicName>
</emphasis>
</heading>
</paragraph>
<paragraph id="8BBC36C52868FFFFF7F0FF59FC42B0C9" blockId="40.[151,1436,222,1147]" box="[151,993,222,247]" pageId="40" pageNumber="41">
<taxonomicName id="4C034D462868FFFFF7F0FF59FE92B0C9" box="[151,305,222,247]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="40" pageNumber="41" phylum="Arthropoda" rank="family">Nymphalidae</taxonomicName>
,
<taxonomicName id="4C034D462868FFFFF65BFF59FE61B0C9" box="[316,450,222,247]" class="Insecta" family="Edelfalter" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="40" pageNumber="41" phylum="Arthropoda" rank="subFamily" subFamily="Libytheinae">Libytheinae</taxonomicName>
(based on similarity rather than on apomorphy).
</paragraph>
</subSubSection>
<subSubSection id="C319654E2868FFFFF7A0FE84FADBB444" pageId="40" pageNumber="41" type="description">
<paragraph id="8BBC36C52868FFFFF7A0FE84FC44B124" blockId="40.[151,1436,222,1147]" box="[199,999,258,283]" pageId="40" pageNumber="41">
<materialsCitation id="3B6B3C982868FFFFF7A0FE84FC44B124" ID-GBIF-Occurrence="1503739244" box="[199,999,258,283]" country="United States of America" county="Teller County" location="Teller County" pageId="40" pageNumber="41" specimenCount="1" stateProvince="Colorado">
<collectingCountry id="F31476552868FFFFF7A0FE84FEA1B125" box="[199,258,259,283]" name="United States of America" pageId="40" pageNumber="41">USA</collectingCountry>
,
<taxonomicName id="4C034D462868FFFFF66AFE85FEDBB125" box="[269,376,258,283]" class="Insecta" family="Papilionidae" genus="Praepapilio" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="40" pageNumber="63" phylum="Arthropoda" rank="species" species="colorado">
<collectingRegion id="49C7F8272868FFFFF66AFE85FEDBB125" box="[269,376,258,283]" country="United States of America" name="Colorado" pageId="40" pageNumber="41">Colorado</collectingRegion>
</taxonomicName>
,
<collectingCounty id="62DD4E492868FFFFF6E4FE85FDBCB125" box="[387,543,258,283]" pageId="40" pageNumber="41">Teller County</collectingCounty>
, Florissant; late Priabonian, late Eocene.
</materialsCitation>
</paragraph>
<paragraph id="8BBC36C52868FFFFF7A0FEA1FD07B101" blockId="40.[151,1436,222,1147]" box="[199,676,294,319]" pageId="40" pageNumber="41">Depository: MCZH (holotype, no. 16353).</paragraph>
<paragraph id="8BBC36C52868FFFFF7A0FECDFE50B1B9" blockId="40.[151,1436,222,1147]" pageId="40" pageNumber="41">
Published figures
<bibRefCitation id="EF924B342868FFFFF6F3FECDFCB8B15D" author="Scudder" box="[404,795,330,355]" pageId="40" pageNumber="41" refString="Scudder, S. H. (1889) The Fossil butterflies of Florissant. United States Geological Survey, 8 th Annual Report, 439 - 472." type="book chapter" year="1889">
Scudder (1889: Pl. LIII
<figureCitation id="13382A402868FFFFF5C2FECDFD57B15D" box="[677,756,330,355]" captionStart="FIGURES 3 4" captionStartId="10.[151,264,1801,1823]" captionTargetBox="[165,1429,840,1774]" captionTargetId="figure@10.[154,1438,828,1788]" captionTargetPageId="10" captionText="FIGURES 3 4. Wing venation of (A) Pareronia Bingham, 1907 (Pieridae), and (B) Hasora Moore, 1881 (Hesperiidae). See text for explanation." httpUri="https://zenodo.org/record/583186/files/figure.png" pageId="40" pageNumber="41">Figs. 4</figureCitation>
9)
</bibRefCitation>
;
<bibRefCitation id="EF924B342868FFFFF44EFECDFBAAB15D" author="Emmel" box="[809,1033,330,355]" pageId="40" pageNumber="41" refString="Emmel, T. C., Minno, M. C. &amp; Drummond, B. A. (1992) Florissant butterflies. A guide to the fossil and present-day species of central Colorado. Stanford University Press, Stanford, x + 118 pp." type="book" year="1992">
Emmel
<emphasis id="B977EAD72868FFFFF4E5FECBFC10B15D" box="[898,947,330,355]" italics="true" pageId="40" pageNumber="41">et al</emphasis>
. (1992
</bibRefCitation>
:
<figureCitation id="13382A402868FFFFF372FECDFBF7B15D" box="[1045,1108,330,355]" 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="40" pageNumber="41">Fig. 1</figureCitation>
/43);
<bibRefCitation id="EF924B342868FFFFF3F2FECCFF1CB1B9" author="Murata" pageId="40" pageNumber="41" 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 44, 45)</bibRefCitation>
;
<bibRefCitation id="EF924B342868FFFFF7ACFEE9FE4CB1B9" author="Kawahara" box="[203,495,366,391]" pageId="40" pageNumber="41" refString="Kawahara, A. Y. (2013) Systematic revision and review of the extant and fossil snout butterflies (Lepidoptera: Nymphalidae: Libytheinae). Zootaxa, 3631 (1), 1 - 74." type="journal article" year="2013">Kawahara (2013: Fig. 63)</bibRefCitation>
.
</paragraph>
<paragraph id="8BBC36C52868FFFFF7A0FE15FBA8B431" blockId="40.[151,1436,222,1147]" pageId="40" pageNumber="41">
The rather poorly preserved specimen shares at least one apomorphic character with
<taxonomicName id="4C034D462868FFFFF3E2FE15FABCB195" box="[1157,1311,402,427]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="40" pageNumber="41" phylum="Arthropoda" rank="family">Nymphalidae</taxonomicName>
, tricarinate antennae. The very long palps are shared with the
<taxonomicName id="4C034D462868FFFFF5B1FE31FCFDB1F1" box="[726,862,438,463]" class="Insecta" family="Edelfalter" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="40" pageNumber="41" phylum="Arthropoda" rank="subFamily" subFamily="Libytheinae">Libytheinae</taxonomicName>
, but are also found in some other nymphalids (see above under
<taxonomicName id="4C034D462868FFFFF656FE5DFD9FB1CD" box="[305,572,474,499]" class="Insecta" family="Edelfalter" genus="Barbarothea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="40" pageNumber="41" phylum="Arthropoda" rank="species" species="florissanti">
<emphasis id="B977EAD72868FFFFF656FE5DFD9FB1CD" box="[305,572,474,499]" italics="true" pageId="40" pageNumber="41">Barbarothea florissanti</emphasis>
</taxonomicName>
). The relatively broad abdomen suggests a female, also indicated by the foreleg bearing complete tarsal claws. The latter is a plesiomorphic character, but in the
<taxonomicName id="4C034D462868FFFFF30DFE79FAA2B229" box="[1130,1281,510,535]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="40" pageNumber="41" phylum="Arthropoda" rank="family">Nymphalidae</taxonomicName>
found only in
<taxonomicName id="4C034D462868FFFFF7F0FDA5FE83B205" box="[151,288,546,571]" class="Insecta" family="Edelfalter" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="40" pageNumber="41" phylum="Arthropoda" rank="subFamily" subFamily="Libytheinae">Libytheinae</taxonomicName>
, a feature in agreement with palpal characters. As far as visible the venation shows plesiomorphic characters, with a radial vein formula 1, 2, 3+(4+5), but large segments of the veins cannot be made out. Hindwing costa apically extended as in recent
<taxonomicName id="4C034D462868FFFFF53FFDEDFCDDB2BD" authority="Fuessly" authorityName="Fuessly" box="[600,894,618,643]" class="Insecta" family="Nymphalidae" genus="Libythea" kingdom="Animalia" order="Lepidoptera" pageId="40" pageNumber="41" phylum="Arthropoda" rank="species" species="celtis">
<emphasis id="B977EAD72868FFFFF53FFDEDFCA7B2BD" box="[600,772,618,643]" italics="true" pageId="40" pageNumber="41">Libythea celtis</emphasis>
(Fuessly)
</taxonomicName>
. Apomorphic character states unique for the
<taxonomicName id="4C034D462868FFFFF7F0FD09FEBEB299" box="[151,285,654,679]" class="Insecta" family="Edelfalter" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="40" pageNumber="41" phylum="Arthropoda" rank="subFamily" subFamily="Libytheinae">Libytheinae</taxonomicName>
are not visible in the fossil, so its assignment to the
<taxonomicName id="4C034D462868FFFFF4F5FD09FBB9B299" box="[914,1050,654,679]" class="Insecta" family="Edelfalter" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="40" pageNumber="41" phylum="Arthropoda" rank="subFamily" subFamily="Libytheinae">Libytheinae</taxonomicName>
, in agreement with
<bibRefCitation id="EF924B342868FFFFF262FD09FF40B2F5" pageId="40" pageNumber="41" refString="Emmel, T. C., Minno, M. C. &amp; Drummond, B. A. (1992) Florissant butterflies. A guide to the fossil and present-day species of central Colorado. Stanford University Press, Stanford, x + 118 pp." type="book">
Emmel
<emphasis id="B977EAD72868FFFFF207FD08FA36B299" box="[1376,1429,654,679]" italics="true" pageId="40" pageNumber="41">et al</emphasis>
. (1992)
</bibRefCitation>
, must be based on similarity only. Scudder emphasized that the fossil is more similar to Old World than to New World
<taxonomicName id="4C034D462868FFFFF641FD51FE13B2D1" box="[294,432,726,751]" class="Insecta" family="Edelfalter" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="40" pageNumber="41" phylum="Arthropoda" rank="subFamily" subFamily="Libytheinae">Libytheinae</taxonomicName>
.
<bibRefCitation id="EF924B342868FFFFF6DAFD51FDDEB2D1" author="Kawahara" box="[445,637,726,751]" pageId="40" pageNumber="41" refString="Kawahara, A. Y. (2009) Phylogeny of snout butterflies (Lepidoptera: Nymphalidae: Libytheinae): combining evidence from the morphology of extant, fossil, and recently extinct taxa. Cladistics, 25, 263 - 278." type="journal article" year="2009">Kawahara (2009</bibRefCitation>
,
<bibRefCitation id="EF924B342868FFFFF5EBFD50FD65B2D1" author="Kawahara" box="[652,710,727,751]" pageId="40" pageNumber="41" refString="Kawahara, A. Y. (2013) Systematic revision and review of the extant and fossil snout butterflies (Lepidoptera: Nymphalidae: Libytheinae). Zootaxa, 3631 (1), 1 - 74." type="journal article" year="2013">2013</bibRefCitation>
) discussed the fossil at length. On the basis of a phylogenetic analysis of all taxa of
<taxonomicName id="4C034D462868FFFFF6F4FD7DFDBFB32D" box="[403,540,762,787]" class="Insecta" family="Edelfalter" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="40" pageNumber="41" phylum="Arthropoda" rank="subFamily" subFamily="Libytheinae">Libytheinae</taxonomicName>
, fossil and recent, he placed the fossil in the extant genus
<taxonomicName id="4C034D462868FFFFF3D4FD7DFA92B32D" box="[1203,1329,762,787]" class="Insecta" family="Nymphalidae" genus="Libytheana" kingdom="Animalia" order="Lepidoptera" pageId="40" pageNumber="41" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD72868FFFFF3D4FD7DFA92B32D" box="[1203,1329,762,787]" italics="true" pageId="40" pageNumber="41">Libytheana</emphasis>
</taxonomicName>
, together with another fossil from the same fossil bed,
<taxonomicName id="4C034D462868FFFFF5F4FC99FC39B309" box="[659,922,798,823]" class="Insecta" family="Edelfalter" genus="Barbarothea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="40" pageNumber="41" phylum="Arthropoda" rank="species" species="florissanti">
<emphasis id="B977EAD72868FFFFF5F4FC99FC39B309" box="[659,922,798,823]" italics="true" pageId="40" pageNumber="41">Barbarothea florissanti</emphasis>
</taxonomicName>
(=
<taxonomicName id="4C034D462868FFFFF4DCFC99FB47B309" box="[955,1252,798,823]" class="Insecta" family="Pieridae" genus="Oligodonta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="40" pageNumber="41" phylum="Arthropoda" rank="species" species="florissantensis">
<emphasis id="B977EAD72868FFFFF4DCFC99FB47B309" box="[955,1252,798,823]" italics="true" pageId="40" pageNumber="41">Oligodonta florissantensis</emphasis>
</taxonomicName>
) (see comments under these names.). The difference between
<taxonomicName id="4C034D462868FFFFF5CAFCC5FC17B365" box="[685,948,834,859]" class="Insecta" family="Nymphalidae" genus="Libytheana" kingdom="Animalia" order="Lepidoptera" pageId="40" pageNumber="41" phylum="Arthropoda" rank="species" species="vagabunda">
<emphasis id="B977EAD72868FFFFF5CAFCC5FC17B365" box="[685,948,834,859]" italics="true" pageId="40" pageNumber="41">Libytheana vagabunda</emphasis>
</taxonomicName>
and
<taxonomicName id="4C034D462868FFFFF494FCC3FB25B365" box="[1011,1158,834,859]" class="Insecta" family="Edelfalter" genus="Barbarothea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="40" pageNumber="41" phylum="Arthropoda" rank="species" species="florissanti">
<emphasis id="B977EAD72868FFFFF494FCC3FB25B365" box="[1011,1158,834,859]" italics="true" pageId="40" pageNumber="41">L. florissanti</emphasis>
</taxonomicName>
is that in the latter the forewing apex is less strongly falcate. Indeed, the relatively strong falcation in
<taxonomicName id="4C034D462868FFFFF36BFCE0FB05B341" box="[1036,1190,870,895]" class="Insecta" family="Nymphalidae" genus="Libytheana" kingdom="Animalia" order="Lepidoptera" pageId="40" pageNumber="41" phylum="Arthropoda" rank="species" species="vagabunda">
<emphasis id="B977EAD72868FFFFF36BFCE0FB05B341" box="[1036,1190,870,895]" italics="true" pageId="40" pageNumber="41">L. vagabunda</emphasis>
</taxonomicName>
is not an observation, but Scudder's reconstruction, indicated by a dashed line. The insects were fossilized ca 34 Ma in a lake environment. The area has been suitable for fossilization of butterflies for a long period, a million to several million years (
<bibRefCitation id="EF924B342868FFFFF783FC55FE16B3D5" author="Smith" box="[228,437,978,1003]" pageId="40" pageNumber="41" refString="Smith, M. E., Carroll, A. R., &amp; Singer, B. S. (2008). Synoptic reconstruction of a major lake system: Eocene Green River Formation, Western United States. Geological Society of America Bulletin, 120, 54 - 84. https: // doi. org / 10.1130 / B 26073.1" type="journal article" year="2008">
Smith
<emphasis id="B977EAD72868FFFFF654FC53FEC4B3D5" box="[307,359,978,1003]" italics="true" pageId="40" pageNumber="41">et al</emphasis>
. 2008
</bibRefCitation>
). Consequently,
<taxonomicName id="4C034D462868FFFFF513FC55FCDAB3D5" box="[628,889,978,1003]" class="Insecta" family="Nymphalidae" genus="Libytheana" kingdom="Animalia" order="Lepidoptera" pageId="40" pageNumber="41" phylum="Arthropoda" rank="species" species="vagabunda">
<emphasis id="B977EAD72868FFFFF513FC55FCDAB3D5" box="[628,889,978,1003]" italics="true" pageId="40" pageNumber="41">Libytheana vagabunda</emphasis>
</taxonomicName>
and
<taxonomicName id="4C034D462868FFFFF4D2FC53FBEBB3D5" box="[949,1096,978,1003]" class="Insecta" family="Edelfalter" genus="Barbarothea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="40" pageNumber="41" phylum="Arthropoda" rank="species" species="florissanti">
<emphasis id="B977EAD72868FFFFF4D2FC53FBEBB3D5" box="[949,1096,978,1003]" italics="true" pageId="40" pageNumber="41">L. florissanti</emphasis>
</taxonomicName>
may have fossilized at times separated long enough for differences to evolve, possibly even at species level.
</paragraph>
<paragraph id="8BBC36C52868FFFFF7A0FB9BFADBB444" blockId="40.[151,1436,222,1147]" pageId="40" pageNumber="41">
<taxonomicName id="4C034D462868FFFFF7A0FB9BFECAB40D" box="[199,361,1050,1075]" class="Insecta" family="Nymphalidae" genus="Libytheana" kingdom="Animalia" order="Lepidoptera" pageId="40" pageNumber="41" phylum="Arthropoda" rank="species" species="vagabunda">
<emphasis id="B977EAD72868FFFFF7A0FB9BFECAB40D" box="[199,361,1050,1075]" italics="true" pageId="40" pageNumber="41">L. vagabunda</emphasis>
</taxonomicName>
as well
<emphasis id="B977EAD72868FFFFF6BAFB9BFD3FB40D" box="[477,668,1050,1075]" italics="true" pageId="40" pageNumber="41">
as
<taxonomicName id="4C034D462868FFFFF562FB9BFD3FB40D" box="[517,668,1050,1075]" class="Insecta" family="Edelfalter" genus="Barbarothea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="40" pageNumber="41" phylum="Arthropoda" rank="species" species="florissanti">L. florissanti</taxonomicName>
</emphasis>
were used as calibration point for the molecular clock of the
<taxonomicName id="4C034D462868FFFFF7F0FBB9FE8DB469" box="[151,302,1086,1111]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="40" pageNumber="41" phylum="Arthropoda" rank="family">Nymphalidae</taxonomicName>
by
<bibRefCitation id="EF924B342868FFFFF63AFBB9FDC7B469" box="[349,612,1086,1111]" pageId="40" pageNumber="41" 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="B977EAD72868FFFFF6B4FBB8FDA4B469" box="[467,519,1086,1111]" italics="true" pageId="40" pageNumber="41">et al</emphasis>
. (2009)
</bibRefCitation>
, the first also employed by Misof
<emphasis id="B977EAD72868FFFFF490FBB8FB88B469" box="[1015,1067,1086,1111]" italics="true" pageId="40" pageNumber="41">et al</emphasis>
. (2014) in their paper on insect evolution for calibrating Macrolepidoptera. See also the section on
<emphasis id="B977EAD72868FFFFF4EEFBE5FAD2B445" box="[905,1393,1122,1147]" italics="true" pageId="40" pageNumber="41">Fossils and calibrating the molecular clock</emphasis>
.
</paragraph>
</subSubSection>
</treatment>
</document>