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<mods:title id="279BE205A87907E6C4814B2BE88236B5">Fossil butterflies, calibration points and the molecular clock (Lepidoptera: Papilionoidea)</mods:title>
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<mods:namePart id="B8ABAA6E93E4689FB036C36F043F33CA">Jong, Rienk De</mods:namePart>
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<treatment id="03AA87D3284EFFD9F7F0FEF7FA3EB445" ID-DOI="http://doi.org/10.5281/zenodo.6046905" ID-GBIF-Taxon="130844900" ID-Zenodo-Dep="6046905" LSID="urn:lsid:plazi:treatment:03AA87D3284EFFD9F7F0FEF7FA3EB445" httpUri="http://treatment.plazi.org/id/03AA87D3284EFFD9F7F0FEF7FA3EB445" lastPageNumber="15" pageId="14" pageNumber="15">
<subSubSection id="C319654E284EFFD9F7F0FEF7FE1AB1F1" pageId="14" pageNumber="15" type="nomenclature">
<paragraph id="8BBC36C5284EFFD9F7F0FEF7FD2BB1B4" blockId="14.[151,648,368,394]" box="[151,648,368,394]" pageId="14" pageNumber="15">
<heading id="D0F481A9284EFFD9F7F0FEF7FD2BB1B4" bold="true" box="[151,648,368,394]" fontSize="11" level="1" pageId="14" pageNumber="15" reason="1">
<emphasis id="B977EAD7284EFFD9F7F0FEF7FD2BB1B4" bold="true" box="[151,648,368,394]" pageId="14" pageNumber="15">
<taxonomicName id="4C034D46284EFFD9F7F0FEF7FF41B1B4" box="[151,226,368,394]" class="Insecta" family="Hesperiidae" genus="Pamphilites" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="species" species="abdita">
<emphasis id="B977EAD7284EFFD9F7F0FEF7FF41B1B4" bold="true" box="[151,226,368,394]" italics="true" pageId="14" pageNumber="15">abdita</emphasis>
</taxonomicName>
.
<taxonomicName id="4C034D46284EFFD9F797FEF7FD2BB1B4" authority="Scudder, 1875" authorityName="Scudder" authorityYear="1875" box="[240,648,368,394]" class="Insecta" family="Hesperiidae" genus="Pamphilites" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="species" species="abdita">
<emphasis id="B977EAD7284EFFD9F797FEF7FE72B1B4" bold="true" box="[240,465,368,394]" italics="true" pageId="14" pageNumber="15">Pamphilites abdita</emphasis>
Scudder, 1875
</taxonomicName>
</emphasis>
</heading>
</paragraph>
<paragraph id="8BBC36C5284EFFD9F7F0FE31FE1AB1F1" blockId="14.[151,1437,438,1147]" box="[151,441,438,463]" pageId="14" pageNumber="15">
<taxonomicName id="4C034D46284EFFD9F7F0FE31FE82B1F1" box="[151,289,438,463]" class="Insecta" family="Hesperiidae" kingdom="Animalia" order="Lepidoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="family">Hesperiidae</taxonomicName>
:
<taxonomicName id="4C034D46284EFFD9F64CFE31FE16B1F1" box="[299,437,438,463]" class="Insecta" family="Hesperiidae" kingdom="Animalia" order="Lepidoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="subFamily" subFamily="Hesperiinae">Hesperiinae</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection id="C319654E284EFFD9F7A0FE5CFA3EB445" pageId="14" pageNumber="15" type="description">
<paragraph id="8BBC36C5284EFFD9F7A0FE5CFA8CB1CD" blockId="14.[151,1437,438,1147]" box="[199,1327,474,499]" pageId="14" pageNumber="15">France, Bouches-du-Rhône, Aix-en-Provence; ChattianAquitanian, Late Oligoceneearly Miocene.</paragraph>
<paragraph id="8BBC36C5284EFFD9F7A0FE79FC48B229" blockId="14.[151,1437,438,1147]" box="[199,1003,510,535]" pageId="14" pageNumber="15">Depositories: MVMF (holotype lost); MNHN (neotype: IPM B-24308).</paragraph>
<paragraph id="8BBC36C5284EFFD9F7A0FDA5FE32B261" blockId="14.[151,1437,438,1147]" pageId="14" pageNumber="15">
Published figures: Scudder (1875: Pl. III, Figs. 14, 17, 18);
<bibRefCitation id="EF924B34284EFFD9F40BFDA4FB64B205" author="Leestmans" box="[876,1223,546,571]" pageId="14" pageNumber="15" refString="Leestmans, R. (1983) Les Lepidopteres fossiles trouves en France (Insecta Lepidoptera). Linneana belgica, 9, 64 - 89." type="journal article" year="1983">
Leestmans (1983:
<figureCitation id="13382A40284EFFD9F35AFDA5FB1DB205" box="[1085,1214,546,571]" captionStart="FIGURES 11 13" captionStartId="34.[151,264,1787,1809]" captionTargetBox="[211,1372,202,1743]" captionTargetId="figure@34.[174,1418,193,1750]" captionTargetPageId="34" captionText="FIGURES 11 13. Ƒanessa pluto Heer, 1849. Figures copied from (1875), Plate II, figures 1, 17 and 7, respectively. (A), copied by Scudder from original description by Heer; (B) and (C), figures sent to Scudder by Mr. Brunner de Wattenwyl, made after the same object. Note differences in wing shape and other details. Figs 14 16. Thaites ruminiana Scudder, 1875. Figures copied from Scudder (1875), Plate III, figures 9, 1 and 3, respectively. (A), underside of fossil; (B), reconstruction of venation; (C), reconstruction of markings. In (A) two separate subcostal veins have been drawn in the forewing, an impossible configuration in Lepidoptera, and apparently a mistake, corrected in (B) and (C). Fig. 17. Cyllo sepulta Boisduval, 1841, wing venation after Scudder (1875). Fig. 18. Butterfly fossil recorded by Wangrin (1939), but shown to be a fraud by Ansorge (2015). Photo kindly provided by J. Ansorge, Horst, Germany (see text for explanation)." httpUri="https://zenodo.org/record/583188/files/figure.png" pageId="14" pageNumber="15">Figs 11, 12</figureCitation>
)
</bibRefCitation>
;
<bibRefCitation id="EF924B34284EFFD9F3B3FDA5FE2FB261" author="Nel" pageId="14" pageNumber="15" refString="Nel, A. &amp; Nel, J. (1986) A propos de deux Lepidopteres fossiles du Stampien d'Aix- en-Provence (Lepidoptera, Hesperiidae et Nymphalidae Satyrinae), Bulletin du Museum National d' Histoire Naturelle, 8, 343 - 350." type="journal article" year="1986">
Nel &amp; Nel (1986:
<figureCitation id="13382A40284EFFD9F7F0FDC1FF77B261" box="[151,212,582,607]" 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="14" pageNumber="15">Fig. 1</figureCitation>
a-b, Pl. I
<figureCitation id="13382A40284EFFD9F627FDC1FE23B261" box="[320,384,582,607]" 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="14" pageNumber="15">Fig. 1</figureCitation>
)
</bibRefCitation>
.
</paragraph>
<paragraph id="8BBC36C5284EFFD9F7A0FDECFC66B2D1" blockId="14.[151,1437,438,1147]" pageId="14" pageNumber="15">
One forewing only. All veins originate from the cell, showing attribution of the fossil to the
<taxonomicName id="4C034D46284EFFD9F383FDEDFACEB2BD" box="[1252,1389,618,643]" class="Insecta" family="Hesperiidae" kingdom="Animalia" order="Lepidoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="family">Hesperiidae</taxonomicName>
; the character recurs in a few subordinate taxa in other superfamilies, but associated with different wing shape. Origin of M2 closer to M3 than to M1, within the
<taxonomicName id="4C034D46284EFFD9F5E8FD35FCB6B2F5" box="[655,789,690,715]" class="Insecta" family="Hesperiidae" kingdom="Animalia" order="Lepidoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="family">Hesperiidae</taxonomicName>
a unique but non-universal character of the
<taxonomicName id="4C034D46284EFFD9F271FD35FA3FB2F5" box="[1302,1436,690,715]" class="Insecta" family="Hesperiidae" kingdom="Animalia" order="Lepidoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="subFamily" subFamily="Hesperiinae">Hesperiinae</taxonomicName>
(
<bibRefCitation id="EF924B34284EFFD9F7F9FD51FED6B2D1" author="Ackery" box="[158,373,726,751]" pageId="14" pageNumber="15" 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="B977EAD7284EFFD9F79DFD50FE89B2D1" box="[250,298,726,751]" italics="true" pageId="14" pageNumber="15">et al</emphasis>
. 1999
</bibRefCitation>
). Wing shape and spotting agree with this subfamily.
</paragraph>
<paragraph id="8BBC36C5284EFFD9F7A0FD7DFD46B3D4" blockId="14.[151,1437,438,1147]" pageId="14" pageNumber="15">
Scudder thought it probable that the fossil wing belonged to a female, apparently only because he found a resemblance with
<taxonomicName id="4C034D46284EFFD9F604FC99FD73B309" authority="Latreille" authorityName="Latreille" box="[355,720,798,823]" class="Insecta" family="Hesperiidae" genus="Pansydia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="species" species="mesogramma">
<emphasis id="B977EAD7284EFFD9F604FC99FDC9B308" box="[355,618,798,823]" italics="true" pageId="14" pageNumber="15">Pansydia mesogramma</emphasis>
Latreille
</taxonomicName>
(now in
<taxonomicName id="4C034D46284EFFD9F451FC99FC11B309" box="[822,946,798,823]" class="Insecta" family="Hesperiidae" genus="Atalopedes" kingdom="Animalia" order="Lepidoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="genus">
<emphasis id="B977EAD7284EFFD9F451FC99FC11B309" box="[822,946,798,823]" italics="true" pageId="14" pageNumber="15">Atalopedes</emphasis>
</taxonomicName>
), a species of the Greater Antilles in which the male has a stigma. The fossil wing lacks any trace of a stigma, but so do the males of many
<taxonomicName id="4C034D46284EFFD9F268FCC5FA3AB365" box="[1295,1433,834,859]" class="Insecta" family="Hesperiidae" kingdom="Animalia" order="Lepidoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="subFamily" subFamily="Hesperiinae">Hesperiinae</taxonomicName>
. Scudder also found a resemblance with
<taxonomicName id="4C034D46284EFFD9F51EFCE0FC1CB341" authority="Fabricius" authorityName="Fabricius" box="[633,959,870,895]" class="Insecta" family="Hesperiidae" genus="Carystus" kingdom="Animalia" order="Lepidoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="species" species="lucasii">
<emphasis id="B977EAD7284EFFD9F51EFCE0FC96B341" box="[633,821,870,895]" italics="true" pageId="14" pageNumber="15">Carystus lucasii</emphasis>
(Fabricius)
</taxonomicName>
, now
<taxonomicName id="4C034D46284EFFD9F36AFCE0FA93B341" authority="Fabricius" authorityName="Fabricius" box="[1037,1328,870,895]" class="Insecta" family="Hesperiidae" genus="Turesis" kingdom="Animalia" order="Lepidoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="species" species="lucas">
<emphasis id="B977EAD7284EFFD9F36AFCE0FB06B341" box="[1037,1189,870,895]" italics="true" pageId="14" pageNumber="15">Turesis lucas</emphasis>
(Fabricius)
</taxonomicName>
, a
<collectingRegion id="49C7F827284EFFD9F23CFCE1FA3FB341" box="[1371,1436,870,895]" country="Tanzania" name="Kusini Unguja" pageId="14" pageNumber="15">South</collectingRegion>
American species. Therefore, he decided a
<collectingRegion id="49C7F827284EFFD9F510FC0DFD1BB39D" box="[631,696,906,931]" country="Tanzania" name="Kusini Unguja" pageId="14" pageNumber="15">South</collectingRegion>
American relationship for this European fossil. Scudder remarked that R4 terminated on the termen close to the apex. I have not seen this condition in any
<taxonomicName id="4C034D46284EFFD9F3F7FC29FABAB3F9" box="[1168,1305,942,967]" class="Insecta" family="Hesperiidae" kingdom="Animalia" order="Lepidoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="family">Hesperiidae</taxonomicName>
, R4 always terminating on the costa just or well before the apex.
</paragraph>
<paragraph id="8BBC36C5284EFFD9F7A0FC71FA3EB445" blockId="14.[151,1437,438,1147]" pageId="14" pageNumber="15">
<materialsCitation id="3B6B3C98284EFFD9F7A0FC71FA3EB445" ID-GBIF-Occurrence="1503739236" county="They" location="New World" municipality="Old World" pageId="14" pageNumber="15" specimenCount="2" stateProvince="South" typeStatus="neotype">
The
<typeStatus id="54B88867284EFFD9F79CFC70FE88B431" box="[251,299,1015,1039]" pageId="14" pageNumber="15">type</typeStatus>
of the fossil species seems to be lost, and
<bibRefCitation id="EF924B34284EFFD9F470FC71FC40B431" author="Nel" box="[791,995,1014,1039]" pageId="14" pageNumber="15" refString="Nel, A. &amp; Nel, J. (1986) A propos de deux Lepidopteres fossiles du Stampien d'Aix- en-Provence (Lepidoptera, Hesperiidae et Nymphalidae Satyrinae), Bulletin du Museum National d' Histoire Naturelle, 8, 343 - 350." type="journal article" year="1986">Nel &amp; Nel (1986)</bibRefCitation>
designated a
<typeStatus id="54B88867284EFFD9F3E6FC70FB79B431" box="[1153,1242,1015,1039]" pageId="14" pageNumber="15" type="neotype">neotype</typeStatus>
that seems to be conspecific with it.
<collectingCounty id="62DD4E49284EFFD9F6EAFB9DFE64B40D" box="[397,455,1050,1075]" pageId="14" pageNumber="15">They</collectingCounty>
correctly remarked that the venation and markings agree with numerous extant
<taxonomicName id="4C034D46284EFFD9F7F0FBB9FEBEB469" box="[151,285,1086,1111]" class="Insecta" family="Hesperiidae" kingdom="Animalia" order="Lepidoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="subFamily" subFamily="Hesperiinae">Hesperiinae</taxonomicName>
species in the
<collectingMunicipality id="6BD8ACBF284EFFD9F6A5FBB9FD9BB469" box="[450,568,1086,1111]" pageId="14" pageNumber="15">Old World</collectingMunicipality>
and
<location id="8EDC601E284EFFD9F508FBB8FD50B469" LSID="urn:lsid:plazi:treatment:03AA87D3284EFFD9F7F0FEF7FA3EB445:8EDC601E284EFFD9F508FBB8FD50B469" box="[623,755,1086,1111]" county="They" municipality="Old World" name="New World" pageId="14" pageNumber="15" stateProvince="South">New World</location>
, so that a
<collectingRegion id="49C7F827284EFFD9F403FBB9FC06B469" box="[868,933,1086,1111]" country="Tanzania" name="Kusini Unguja" pageId="14" pageNumber="15">South</collectingRegion>
American affinity of the fossil species is not obvious.
<location id="8EDC601E284EFFD9F79AFBE5FEB3B445" LSID="urn:lsid:plazi:treatment:03AA87D3284EFFD9F7F0FEF7FA3EB445:8EDC601E284EFFD9F79AFBE5FEB3B445" box="[253,272,1122,1147]" county="They" municipality="Old World" name="If" pageId="14" pageNumber="15" stateProvince="South">If</location>
used for calibration, the fossil should be used as an estimated minimum age for the root of
<taxonomicName id="4C034D46284EFFD9F268FBE5FA3AB445" box="[1295,1433,1122,1147]" class="Insecta" family="Hesperiidae" kingdom="Animalia" order="Lepidoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="subFamily" subFamily="Hesperiinae">Hesperiinae</taxonomicName>
.
</materialsCitation>
</paragraph>
</subSubSection>
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