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<document id="1CE5679AB64C64560000CBFE5A9B8889" ID-DOI="10.5281/zenodo.188448" ID-GBIF-Dataset="10d3b521-818c-44d2-af1b-df4281f82b80" ID-ISSN="1175-5326" ID-Zenodo-Dep="188448" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1461093309137" checkinUser="plazi" docAuthor="Garzón-Orduña, Ivonne J. &amp; Penz, Carla M." docDate="2009" docId="4F1326670D26755D6BD0B7E76694A4F1" docLanguage="en" docName="zt02134p022.pdf" docOrigin="Zootaxa 2134" docStyle="DocumentStyle:890A69B780ED73D6DB8551B71C8AC79E.4:Zootaxa.2009-2012.journal_article" docStyleId="890A69B780ED73D6DB8551B71C8AC79E" docStyleName="Zootaxa.2009-2012.journal_article" docStyleVersion="4" docTitle="Brassolis astyra Godart" docType="treatment" docVersion="9" lastPageNumber="10" masterDocId="B32A5E1F0D2175546B47B2416662A72F" masterDocTitle="Phylogeny of Dynastor and Brassolis butterflies (Lepidoptera: Nymphalidae): a tough nut to crack" masterLastPageNumber="22" masterPageNumber="1" pageNumber="8" updateTime="1698588535132" updateUser="plazi">
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<mods:title id="29B4D83000E45C2F08A4EEE67ABCF9CB">Phylogeny of Dynastor and Brassolis butterflies (Lepidoptera: Nymphalidae): a tough nut to crack</mods:title>
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<mods:namePart id="EB038F90A17BD747582BF8EF897ECFCF">Garzón-Orduña, Ivonne J.</mods:namePart>
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<mods:namePart id="4A0E3F59253747D24DE5846A1400F7EA">Penz, Carla M.</mods:namePart>
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<mods:date id="54512D1562026480434C10DD51ACCB12">2009</mods:date>
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<treatment id="4F1326670D26755D6BD0B7E76694A4F1" ID-DOI="http://doi.org/10.5281/zenodo.6221631" ID-GBIF-Taxon="119619938" ID-Zenodo-Dep="6221631" LSID="urn:lsid:plazi:treatment:4F1326670D26755D6BD0B7E76694A4F1" httpUri="http://treatment.plazi.org/id/4F1326670D26755D6BD0B7E76694A4F1" lastPageId="9" lastPageNumber="10" pageId="7" pageNumber="8">
<subSubSection id="8FA0C4FA0D2675536BD0B7E767C0A2CC" pageId="7" pageNumber="8" type="nomenclature">
<paragraph id="C70597710D2675536BD0B7E767DAA2EF" blockId="7.[151,440,1446,1507]" box="[151,440,1446,1472]" pageId="7" pageNumber="8">
<heading id="9C4D201D0D2675536BD0B7E767DAA2EF" bold="true" box="[151,440,1446,1472]" fontSize="11" level="1" pageId="7" pageNumber="8" reason="1">
<taxonomicName id="00BAECF20D2675536BD0B7E767DAA2EF" ID-CoL="N86Q" authority="Godart" authorityName="Godart" box="[151,440,1446,1472]" class="Insecta" family="Nymphalidae" genus="Brassolis" kingdom="Animalia" order="Lepidoptera" pageId="7" pageNumber="8" phylum="Arthropoda" rank="species" species="astyra">
<emphasis id="F5CE4B630D2675536BD0B7E767DAA2EF" bold="true" box="[151,440,1446,1472]" pageId="7" pageNumber="8">
<emphasis id="F5CE4B630D2675536BD0B7E76734A2EF" bold="true" box="[151,342,1446,1472]" italics="true" pageId="7" pageNumber="8">Brassolis astyra</emphasis>
Godart
</emphasis>
</taxonomicName>
</heading>
</paragraph>
<paragraph id="C70597710D2675536BD0B78867C0A2CC" blockId="7.[151,440,1446,1507]" box="[151,418,1481,1507]" pageId="7" pageNumber="8">
(
<figureCitation id="5F818BF40D2675536BD8B7886681A2CC" box="[159,227,1481,1507]" captionStart="FIGURE 3" captionStartId="6.[151,257,1913,1937]" captionTargetBox="[223,1316,798,1878]" captionTargetId="figure@6.[218,1348,783,1910]" captionTargetPageId="6" captionText="FIGURE 3. Brassolis species, in dorsal (left) and ventral (right) views (except for C). (A &amp; B) Male and female B. isthmia from Panama. (C) Male B. isthmia from Costa Rica. (D) Male of granadensis from Colombia. (E) Detail showing head and proboscis of B. isthmia from Panama. (F &amp; G) Male and female B. astyra from Rio de Janeiro (Brazil). (H) Male B. astyra from Rio Grande do Sul (Brazil)." httpUri="https://zenodo.org/record/188451/files/figure.png" pageId="7" pageNumber="8">Fig. 3</figureCitation>
FH; 8D, I; 9D)
</paragraph>
</subSubSection>
<subSubSection id="8FA0C4FA0D2675536BD0B45367ECA04D" pageId="7" pageNumber="8" type="diagnosis">
<paragraph id="C70597710D2675536BD0B45367ECA04D" blockId="7.[151,1437,1554,2002]" pageId="7" pageNumber="8">
<emphasis id="F5CE4B630D2675536BD0B4536778A103" bold="true" box="[151,282,1554,1580]" pageId="7" pageNumber="8">Diagnosis:</emphasis>
Recognized by a combination of five characters: (1) in both sexes, dorsal FW postmedial orange band clearly bifurcated anteriorly (above M3). In females this band is faded above M3 and inside the discal cell; (2) specimens from the northern end of the distribution (
<taxonomicName id="00BAECF20D26755368C0B421624AA156" box="[903,1064,1632,1657]" class="Insecta" family="Nymphalidae" genus="Brassolis" kingdom="Animalia" order="Lepidoptera" pageId="7" pageNumber="8" phylum="Arthropoda" rank="subSpecies" species="astyra" subSpecies="astyra">
<emphasis id="F5CE4B630D26755368C0B421624AA156" box="[903,1064,1632,1657]" italics="true" pageId="7" pageNumber="8">astyra astyra</emphasis>
</taxonomicName>
) lack a dorsal HW postmedial band, but this band varies from faded to well developed in specimens from southern localities (
<taxonomicName id="00BAECF20D2675536E09B4C66764A1E9" class="Insecta" family="Nymphalidae" genus="Brassolis" kingdom="Animalia" order="Lepidoptera" pageId="7" pageNumber="8" phylum="Arthropoda" rank="subSpecies" species="astyra" subSpecies="philocala">
<emphasis id="F5CE4B630D2675536E09B4C66764A1E9" italics="true" pageId="7" pageNumber="8">astyra philocala</emphasis>
</taxonomicName>
). Dorsal HW postmedial band generally narrower and more jagged in
<taxonomicName id="00BAECF20D2675536F04B4EC6341A1E9" box="[1091,1315,1709,1734]" class="Insecta" family="Nymphalidae" genus="Brassolis" kingdom="Animalia" order="Lepidoptera" pageId="7" pageNumber="8" phylum="Arthropoda" rank="subSpecies" species="astyra" subSpecies="philocala">
<emphasis id="F5CE4B630D2675536F04B4EC6341A1E9" box="[1091,1315,1709,1734]" italics="true" pageId="7" pageNumber="8">B. astyra philocala</emphasis>
</taxonomicName>
than in
<taxonomicName id="00BAECF20D2675536EC4B4EC6769A1C2" class="Insecta" family="Nymphalidae" genus="Brassolis" kingdom="Animalia" order="Lepidoptera" pageId="7" pageNumber="8" phylum="Arthropoda" rank="species" species="sophorae">
<emphasis id="F5CE4B630D2675536EC4B4EC6769A1C2" italics="true" pageId="7" pageNumber="8">B. sophorae</emphasis>
</taxonomicName>
and
<taxonomicName id="00BAECF20D2675536A0EB49567D1A1C2" box="[329,435,1748,1773]" class="Insecta" family="Nymphalidae" genus="Brassolis" kingdom="Animalia" order="Lepidoptera" pageId="7" pageNumber="8" phylum="Arthropoda" rank="species" species="dinizi">
<emphasis id="F5CE4B630D2675536A0EB49567D1A1C2" box="[329,435,1748,1773]" italics="true" pageId="7" pageNumber="8">B. dinizi</emphasis>
</taxonomicName>
; (3) in both sexes, abdomen orange dorsally from third segment onward; (4) in posterior view, male valva usually with a small point at apex (
<figureCitation id="5F818BF40D267553682AB4BB65D0A03B" box="[877,946,1786,1812]" captionStart="FIGURE 8" captionStartId="14.[151,255,1625,1649]" captionTargetBox="[157,1409,196,1582]" captionTargetId="figure@14.[151,1436,164,1735]" captionTargetPageId="14" captionText="FIGURE 8. Male characters. Scale bars, 1 mm. Labial palp in lateral view. (A) Bia actorion. (B) Brassolis sophorae. (C) Dynastor darius. Tegumen in dorsal view. (D) B. astyra. Genitalic capsule in lateral view. (E) D. darius. (F) B. sophorae. Tegumen and uncus in lateral view. (G) D. macrosiris. Phallus in lateral view. (H) D. napoleon. Juxta in ventral view. (I) B. astyra. (J) D. napoleon. Phallus in ventral view. (K) D. darius. (L) B. haenschi. (M) B. granadensis." httpUri="https://zenodo.org/record/188456/files/figure.png" pageId="7" pageNumber="8">Fig. 8</figureCitation>
D); (5) in ventral view, anterior section of the female sterigma interrupted at mid point, sclerotized loop originates from a fold closer to the midline than in other species.
</paragraph>
</subSubSection>
<subSubSection id="8FA0C4FA0D26755D6B82B52F6694A4F1" lastPageId="9" lastPageNumber="10" pageId="7" pageNumber="8" type="distribution">
<paragraph id="C70597710D2675536B82B52F6520A0A7" blockId="7.[151,1437,1554,2002]" box="[197,834,1902,1928]" pageId="7" pageNumber="8">
<emphasis id="F5CE4B630D2675536B82B52F670AA0A7" bold="true" box="[197,360,1902,1928]" pageId="7" pageNumber="8">Distribution:</emphasis>
Southeastern
<collectingCountry id="BFADD7E10D267553694AB52F6437A0A7" box="[525,597,1902,1928]" name="Brazil" pageId="7" pageNumber="8">Brazil</collectingCountry>
(
<bibRefCitation id="A32BEA800D2675536923B52F6554A0A7" author="Casagrande" box="[612,822,1902,1928]" pageId="7" pageNumber="17" refString="Casagrande, M. M. (2004) Brassolini. Atlas of Neotropical Lepidoptera (ed. By J. B. Heppner), Part 4 A Checklist (ed. By G. Lamas), pp. 201 - 205. Association of Tropical Lepidoptera / Scientific Publishers, Gainesville, FL." type="book chapter" year="2004">Casagrande 2004</bibRefCitation>
).
</paragraph>
<paragraph id="C70597710D2675536BD0B5D46459A0FD" blockId="7.[151,1437,1554,2002]" pageId="7" pageNumber="8">
<emphasis id="F5CE4B630D2675536BD0B5D4674AA080" bold="true" box="[151,296,1941,1967]" pageId="7" pageNumber="8">Subspecies:</emphasis>
<bibRefCitation id="A32BEA800D2675536A68B5D4646FA080" author="Casagrande" box="[303,525,1941,1967]" pageId="7" pageNumber="17" refString="Casagrande, M. M. (2004) Brassolini. Atlas of Neotropical Lepidoptera (ed. By J. B. Heppner), Part 4 A Checklist (ed. By G. Lamas), pp. 201 - 205. Association of Tropical Lepidoptera / Scientific Publishers, Gainesville, FL." type="book chapter" year="2004">Casagrande (2004)</bibRefCitation>
listed two subspecies: nominal
<taxonomicName id="00BAECF20D26755368C4B5D465AEA081" box="[899,972,1941,1966]" class="Insecta" family="Nymphalidae" genus="Brassolis" kingdom="Animalia" order="Lepidoptera" pageId="7" pageNumber="8" phylum="Arthropoda" rank="species" species="astyra">
<emphasis id="F5CE4B630D26755368C4B5D465AEA081" box="[899,972,1941,1966]" italics="true" pageId="7" pageNumber="8">astyra</emphasis>
</taxonomicName>
(
<typeStatus id="180129D30D267553689CB5D4626DA080" box="[987,1039,1941,1967]" pageId="7" pageNumber="8">type</typeStatus>
locality
<collectingCountry id="BFADD7E10D2675536F31B5D462A0A080" box="[1142,1218,1941,1967]" name="Brazil" pageId="7" pageNumber="8">Brazil</collectingCountry>
), and
<emphasis id="F5CE4B630D2675536E4FB5D46315A081" box="[1288,1399,1941,1966]" italics="true" pageId="7" pageNumber="8">philocala</emphasis>
Stichel (
<collectingCountry id="BFADD7E10D2675536BB0B5F96721A0FD" box="[247,323,1976,2002]" name="Brazil" pageId="7" pageNumber="8">Brazil</collectingCountry>
, Rio Grande do Sul).
</paragraph>
<paragraph id="C70597710D29755C6BD0B2D96706A79D" blockId="8.[151,1437,152,2034]" box="[151,356,152,178]" pageId="8" pageNumber="9">Cladistic analysis</paragraph>
<paragraph id="C70597710D29755C6B82B2FF64E3A6C8" blockId="8.[151,1437,152,2034]" pageId="8" pageNumber="9">
Implied weights searches under K=1 and 2 found two most parsimonious trees. Higher values of K (K=3 to 20) consistently yielded the same three most parsimonious trees, one of which is the topology of the strict consensus of these three trees (
<figureCitation id="5F818BF40D29755C6AB8B34D6427A609" box="[511,581,268,294]" captionStart="FIGURE 4" captionStartId="10.[151,261,1680,1704]" captionTargetBox="[207,1385,425,1637]" captionTargetId="figure@10.[165,1425,391,1651]" captionTargetPageId="10" captionText="FIGURE 4. (A) Strict consensus tree from the analysis of 57 characters with K = 3. (B &amp; C) Two of the most parsimonious trees, the third tree had the same topology of the strict consensus. Relative Bremer support (on the left) and Jackknife values (on the right) are on the left hand of the branches." httpUri="https://zenodo.org/record/188452/files/figure.png" pageId="8" pageNumber="9">Fig. 4</figureCitation>
A). The other two are the same trees found under K=1 and 2 (
<figureCitation id="5F818BF40D29755C6E57B34D6335A609" box="[1296,1367,268,294]" captionStart="FIGURE 4" captionStartId="10.[151,261,1680,1704]" captionTargetBox="[207,1385,425,1637]" captionTargetId="figure@10.[165,1425,391,1651]" captionTargetPageId="10" captionText="FIGURE 4. (A) Strict consensus tree from the analysis of 57 characters with K = 3. (B &amp; C) Two of the most parsimonious trees, the third tree had the same topology of the strict consensus. Relative Bremer support (on the left) and Jackknife values (on the right) are on the left hand of the branches." httpUri="https://zenodo.org/record/188452/files/figure.png" pageId="8" pageNumber="9">Fig. 4</figureCitation>
B, C). Trees had a fit of 5.2. The three most parsimonious trees show
<taxonomicName id="00BAECF20D29755C683CB373658AA664" box="[891,1000,306,331]" class="Insecta" family="Nymphalidae" genus="Dynastor" kingdom="Animalia" order="Lepidoptera" pageId="8" pageNumber="9" phylum="Arthropoda" rank="genus">
<emphasis id="F5CE4B630D29755C683CB373658AA664" box="[891,1000,306,331]" italics="true" pageId="8" pageNumber="9">Dynastor</emphasis>
</taxonomicName>
and
<taxonomicName id="00BAECF20D29755C6F66B37362ECA664" box="[1057,1166,306,331]" class="Insecta" family="Nymphalidae" genus="Brassolis" kingdom="Animalia" order="Lepidoptera" pageId="8" pageNumber="9" phylum="Arthropoda" rank="genus">
<emphasis id="F5CE4B630D29755C6F66B37362ECA664" box="[1057,1166,306,331]" italics="true" pageId="8" pageNumber="9">Brassolis</emphasis>
</taxonomicName>
as monophyletic sister genera, but differed in the relative positions of
<emphasis id="F5CE4B630D29755C6986B3186379A65D" box="[705,1307,345,370]" italics="true" pageId="8" pageNumber="9">
<taxonomicName id="00BAECF20D29755C6986B3186530A65D" box="[705,850,345,370]" class="Insecta" family="Nymphalidae" genus="Brassolis" kingdom="Animalia" order="Lepidoptera" pageId="8" pageNumber="9" phylum="Arthropoda" rank="species" species="sophorae">B. sophorae</taxonomicName>
,
<taxonomicName id="00BAECF20D29755C6819B3186588A65D" box="[862,1002,345,370]" class="Insecta" family="Nymphalidae" genus="Brassolis" kingdom="Animalia" order="Lepidoptera" pageId="8" pageNumber="9" phylum="Arthropoda" rank="species" species="haenschi">B. haenschi</taxonomicName>
,
<taxonomicName id="00BAECF20D29755C68B0B318623EA65D" box="[1015,1116,345,370]" class="Insecta" family="Nymphalidae" genus="Brassolis" kingdom="Animalia" order="Lepidoptera" pageId="8" pageNumber="9" phylum="Arthropoda" rank="species" species="dinizi">B. dinizi</taxonomicName>
, and
<taxonomicName id="00BAECF20D29755C6FDBB3186374A65D" box="[1180,1302,345,370]" class="Insecta" family="Nymphalidae" genus="Brassolis" kingdom="Animalia" order="Lepidoptera" pageId="8" pageNumber="9" phylum="Arthropoda" rank="species" species="isthmia">B. isthmia</taxonomicName>
,
</emphasis>
which had an effect on the resolution of the strict consensus (compare topologies in
<figureCitation id="5F818BF40D29755C6F5EB3C16206A6B5" box="[1049,1124,384,410]" captionStart="FIGURE 4" captionStartId="10.[151,261,1680,1704]" captionTargetBox="[207,1385,425,1637]" captionTargetId="figure@10.[165,1425,391,1651]" captionTargetPageId="10" captionText="FIGURE 4. (A) Strict consensus tree from the analysis of 57 characters with K = 3. (B &amp; C) Two of the most parsimonious trees, the third tree had the same topology of the strict consensus. Relative Bremer support (on the left) and Jackknife values (on the right) are on the left hand of the branches." httpUri="https://zenodo.org/record/188452/files/figure.png" pageId="8" pageNumber="9">Fig. 4</figureCitation>
). Unambiguous character transformations for the nodes and terminals were optimized in the strict consensus tree using “hard” polytomy option and are described below (
<figureCitation id="5F818BF40D29755C6952B38C643EA6C8" box="[533,604,461,487]" captionStart="FIGURE 4" captionStartId="10.[151,261,1680,1704]" captionTargetBox="[207,1385,425,1637]" captionTargetId="figure@10.[165,1425,391,1651]" captionTargetPageId="10" captionText="FIGURE 4. (A) Strict consensus tree from the analysis of 57 characters with K = 3. (B &amp; C) Two of the most parsimonious trees, the third tree had the same topology of the strict consensus. Relative Bremer support (on the left) and Jackknife values (on the right) are on the left hand of the branches." httpUri="https://zenodo.org/record/188452/files/figure.png" pageId="8" pageNumber="9">Fig. 4</figureCitation>
A).
</paragraph>
<paragraph id="C70597710D29755C6B82B3B5627EA5E0" blockId="8.[151,1437,152,2034]" pageId="8" pageNumber="9">
Four synapomorphies supported the
<taxonomicName id="00BAECF20D29755C6937B3B564BFA522" box="[624,733,500,525]" class="Insecta" family="Nymphalidae" genus="Dynastor" kingdom="Animalia" order="Lepidoptera" pageId="8" pageNumber="9" phylum="Arthropoda" rank="genus">
<emphasis id="F5CE4B630D29755C6937B3B564BFA522" box="[624,733,500,525]" italics="true" pageId="8" pageNumber="9">Dynastor</emphasis>
</taxonomicName>
+
<taxonomicName id="00BAECF20D29755C69BBB3B5650BA522" box="[764,873,500,525]" class="Insecta" family="Nymphalidae" genus="Brassolis" kingdom="Animalia" order="Lepidoptera" pageId="8" pageNumber="9" phylum="Arthropoda" rank="genus">
<emphasis id="F5CE4B630D29755C69BBB3B5650BA522" box="[764,873,500,525]" italics="true" pageId="8" pageNumber="9">Brassolis</emphasis>
</taxonomicName>
clade: last segment of labial palpus shorter than the distal width of second segment (character 1:2,
<figureCitation id="5F818BF40D29755C69AEB05B654DA51B" box="[745,815,538,564]" captionStart="FIGURE 8" captionStartId="14.[151,255,1625,1649]" captionTargetBox="[157,1409,196,1582]" captionTargetId="figure@14.[151,1436,164,1735]" captionTargetPageId="14" captionText="FIGURE 8. Male characters. Scale bars, 1 mm. Labial palp in lateral view. (A) Bia actorion. (B) Brassolis sophorae. (C) Dynastor darius. Tegumen in dorsal view. (D) B. astyra. Genitalic capsule in lateral view. (E) D. darius. (F) B. sophorae. Tegumen and uncus in lateral view. (G) D. macrosiris. Phallus in lateral view. (H) D. napoleon. Juxta in ventral view. (I) B. astyra. (J) D. napoleon. Phallus in ventral view. (K) D. darius. (L) B. haenschi. (M) B. granadensis." httpUri="https://zenodo.org/record/188456/files/figure.png" pageId="8" pageNumber="9">Fig. 8</figureCitation>
B), reversed to state
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<taxonomicName id="00BAECF20D29755C6F1FB05B6280A51C" box="[1112,1250,538,563]" class="Insecta" family="Nymphalidae" genus="Brassolis" kingdom="Animalia" order="Lepidoptera" pageId="8" pageNumber="9" phylum="Arthropoda" rank="species" species="haenschi">
<emphasis id="F5CE4B630D29755C6F1FB05B6280A51C" box="[1112,1250,538,563]" italics="true" pageId="8" pageNumber="9">B. haenschi</emphasis>
</taxonomicName>
(approximately the same length as distal width of second segment); valva not projecting beyond uncus tip (30:1,
<figureCitation id="5F818BF40D29755C6E76B000631AA574" box="[1329,1400,577,603]" captionStart="FIGURE 8" captionStartId="14.[151,255,1625,1649]" captionTargetBox="[157,1409,196,1582]" captionTargetId="figure@14.[151,1436,164,1735]" captionTargetPageId="14" captionText="FIGURE 8. Male characters. Scale bars, 1 mm. Labial palp in lateral view. (A) Bia actorion. (B) Brassolis sophorae. (C) Dynastor darius. Tegumen in dorsal view. (D) B. astyra. Genitalic capsule in lateral view. (E) D. darius. (F) B. sophorae. Tegumen and uncus in lateral view. (G) D. macrosiris. Phallus in lateral view. (H) D. napoleon. Juxta in ventral view. (I) B. astyra. (J) D. napoleon. Phallus in ventral view. (K) D. darius. (L) B. haenschi. (M) B. granadensis." httpUri="https://zenodo.org/record/188456/files/figure.png" pageId="8" pageNumber="9">Fig. 8</figureCitation>
E); groove at the articulating point between valva and appendices angulares limited to the base of valva (34:1); distal opening of phallus located dorsolaterally (38:1).
<taxonomicName id="00BAECF20D29755C6806B0CE65D1A587" box="[833,947,655,680]" class="Insecta" family="Nymphalidae" genus="Dynastor" kingdom="Animalia" order="Lepidoptera" pageId="8" pageNumber="9" phylum="Arthropoda" rank="genus">
<emphasis id="F5CE4B630D29755C6806B0CE65D1A587" box="[833,947,655,680]" italics="true" pageId="8" pageNumber="9">Dynastor</emphasis>
</taxonomicName>
+
<taxonomicName id="00BAECF20D29755C6892B0CE6225A587" box="[981,1095,655,680]" class="Insecta" family="Nymphalidae" genus="Brassolis" kingdom="Animalia" order="Lepidoptera" pageId="8" pageNumber="9" phylum="Arthropoda" rank="genus">
<emphasis id="F5CE4B630D29755C6892B0CE6225A587" box="[981,1095,655,680]" italics="true" pageId="8" pageNumber="9">Brassolis</emphasis>
</taxonomicName>
had a low Relative Bremer support and a moderate value of jackknife recuperation (30 and 59;
<figureCitation id="5F818BF40D29755C68F6B0F46595A5E0" box="[945,1015,693,719]" captionStart="FIGURE 4" captionStartId="10.[151,261,1680,1704]" captionTargetBox="[207,1385,425,1637]" captionTargetId="figure@10.[165,1425,391,1651]" captionTargetPageId="10" captionText="FIGURE 4. (A) Strict consensus tree from the analysis of 57 characters with K = 3. (B &amp; C) Two of the most parsimonious trees, the third tree had the same topology of the strict consensus. Relative Bremer support (on the left) and Jackknife values (on the right) are on the left hand of the branches." httpUri="https://zenodo.org/record/188452/files/figure.png" pageId="8" pageNumber="9">Fig. 4</figureCitation>
A).
</paragraph>
<paragraph id="C70597710D29755C6B82B09D65C0A32B" blockId="8.[151,1437,152,2034]" pageId="8" pageNumber="9">
The monophyly of the genus
<taxonomicName id="00BAECF20D29755C6960B09D64F4A5DA" box="[551,662,732,757]" class="Insecta" family="Nymphalidae" genus="Dynastor" kingdom="Animalia" order="Lepidoptera" pageId="8" pageNumber="9" phylum="Arthropoda" rank="genus">
<emphasis id="F5CE4B630D29755C6960B09D64F4A5DA" box="[551,662,732,757]" italics="true" pageId="8" pageNumber="9">Dynastor</emphasis>
</taxonomicName>
was supported by seven synapomorphies and one homoplasious transformation. The synapomorphies were: dorsal forewing costal margin striped (4:1,
<figureCitation id="5F818BF40D29755C6F91B1436342A433" box="[1238,1312,770,796]" captionStart="FIGURE 7" captionStartId="13.[151,255,1907,1931]" captionTargetBox="[202,1408,195,1834]" captionTargetId="figure@13.[151,1436,157,1880]" captionTargetPageId="13" captionText="FIGURE 7. Wing characters. (A) Brassolis dizini. (B) B. astyra. (C) B. haenschi. (D) Dynastor darius." httpUri="https://zenodo.org/record/188455/files/figure.png" pageId="8" pageNumber="9">Fig. 7</figureCitation>
D), dorsal forewing postmedial band broken (8:1,
<figureCitation id="5F818BF40D29755C692DB16864D2A46C" box="[618,688,809,835]" captionStart="FIGURE 7" captionStartId="13.[151,255,1907,1931]" captionTargetBox="[202,1408,195,1834]" captionTargetId="figure@13.[151,1436,157,1880]" captionTargetPageId="13" captionText="FIGURE 7. Wing characters. (A) Brassolis dizini. (B) B. astyra. (C) B. haenschi. (D) Dynastor darius." httpUri="https://zenodo.org/record/188455/files/figure.png" pageId="8" pageNumber="9">Fig. 7</figureCitation>
D), female foreleg distal tarsomere with ventral spines between the rows of sensilla (24:1,
<figureCitation id="5F818BF40D29755C6A97B1116477A445" box="[464,533,848,874]" captionStart="FIGURE 9" captionStartId="15.[151,256,1456,1480]" captionTargetBox="[153,1412,201,1455]" captionTargetId="figure@15.[151,1436,127,1497]" captionTargetPageId="15" captionText="FIGURE 9. Female characters. Scale bars, 1 mm. Female distal tarsomere in ventral view (details of pulvillar process omitted). (A) Dynastor darius. Sterigma in ventral view. (B) D. darius. (C) B. isthmia. (D) B. astyra. (E) B. dinizi. Posterior lobes of papillae annales in lateral view. (F) Elymnias hypermnestra. (G) D. napoleon. (H) B. haenschi. Note that the lamella antevaginalis is not visible in C, D, and E due to the orientation of the drawing." httpUri="https://zenodo.org/record/188457/files/figure.png" pageId="8" pageNumber="9">Fig. 9</figureCitation>
A), dorsal edge of juxta continuous (37:1), sclerotized tip of phallus forming a small coecum that projects beyond the distal opening (44:1,
<figureCitation id="5F818BF40D29755C682AB13765D2A4BF" box="[877,944,886,912]" captionStart="FIGURE 8" captionStartId="14.[151,255,1625,1649]" captionTargetBox="[157,1409,196,1582]" captionTargetId="figure@14.[151,1436,164,1735]" captionTargetPageId="14" captionText="FIGURE 8. Male characters. Scale bars, 1 mm. Labial palp in lateral view. (A) Bia actorion. (B) Brassolis sophorae. (C) Dynastor darius. Tegumen in dorsal view. (D) B. astyra. Genitalic capsule in lateral view. (E) D. darius. (F) B. sophorae. Tegumen and uncus in lateral view. (G) D. macrosiris. Phallus in lateral view. (H) D. napoleon. Juxta in ventral view. (I) B. astyra. (J) D. napoleon. Phallus in ventral view. (K) D. darius. (L) B. haenschi. (M) B. granadensis." httpUri="https://zenodo.org/record/188456/files/figure.png" pageId="8" pageNumber="9">Fig. 8</figureCitation>
H), lateral uncus wings merging to form a single dorsal keel (47:1) and distal portion of uncus tall (48:1,
<figureCitation id="5F818BF40D29755C682BB1DC65CDA498" box="[876,943,925,951]" captionStart="FIGURE 8" captionStartId="14.[151,255,1625,1649]" captionTargetBox="[157,1409,196,1582]" captionTargetId="figure@14.[151,1436,164,1735]" captionTargetPageId="14" captionText="FIGURE 8. Male characters. Scale bars, 1 mm. Labial palp in lateral view. (A) Bia actorion. (B) Brassolis sophorae. (C) Dynastor darius. Tegumen in dorsal view. (D) B. astyra. Genitalic capsule in lateral view. (E) D. darius. (F) B. sophorae. Tegumen and uncus in lateral view. (G) D. macrosiris. Phallus in lateral view. (H) D. napoleon. Juxta in ventral view. (I) B. astyra. (J) D. napoleon. Phallus in ventral view. (K) D. darius. (L) B. haenschi. (M) B. granadensis." httpUri="https://zenodo.org/record/188456/files/figure.png" pageId="8" pageNumber="9">Fig. 8</figureCitation>
G). The homoplasious transformation was the ventral outline of phallobase evenly arched (45:0,
<figureCitation id="5F818BF40D29755C687BB18565E7A4F1" box="[828,901,964,990]" captionStart="FIGURE 8" captionStartId="14.[151,255,1625,1649]" captionTargetBox="[157,1409,196,1582]" captionTargetId="figure@14.[151,1436,164,1735]" captionTargetPageId="14" captionText="FIGURE 8. Male characters. Scale bars, 1 mm. Labial palp in lateral view. (A) Bia actorion. (B) Brassolis sophorae. (C) Dynastor darius. Tegumen in dorsal view. (D) B. astyra. Genitalic capsule in lateral view. (E) D. darius. (F) B. sophorae. Tegumen and uncus in lateral view. (G) D. macrosiris. Phallus in lateral view. (H) D. napoleon. Juxta in ventral view. (I) B. astyra. (J) D. napoleon. Phallus in ventral view. (K) D. darius. (L) B. haenschi. (M) B. granadensis." httpUri="https://zenodo.org/record/188456/files/figure.png" pageId="8" pageNumber="9">Fig. 8</figureCitation>
H). Relative Bremer support and Jackknife recuperation values for
<taxonomicName id="00BAECF20D29755C6AEBB1AB647BA32C" box="[428,537,1002,1027]" class="Insecta" family="Nymphalidae" genus="Dynastor" kingdom="Animalia" order="Lepidoptera" pageId="8" pageNumber="9" phylum="Arthropoda" rank="genus">
<emphasis id="F5CE4B630D29755C6AEBB1AB647BA32C" box="[428,537,1002,1027]" italics="true" pageId="8" pageNumber="9">Dynastor</emphasis>
</taxonomicName>
were high (100 and 99;
<figureCitation id="5F818BF40D29755C6870B1AB651FA32B" box="[823,893,1002,1028]" captionStart="FIGURE 4" captionStartId="10.[151,261,1680,1704]" captionTargetBox="[207,1385,425,1637]" captionTargetId="figure@10.[165,1425,391,1651]" captionTargetPageId="10" captionText="FIGURE 4. (A) Strict consensus tree from the analysis of 57 characters with K = 3. (B &amp; C) Two of the most parsimonious trees, the third tree had the same topology of the strict consensus. Relative Bremer support (on the left) and Jackknife values (on the right) are on the left hand of the branches." httpUri="https://zenodo.org/record/188452/files/figure.png" pageId="8" pageNumber="9">Fig. 4</figureCitation>
A).
</paragraph>
<paragraph id="C70597710D29755C6B82B6506763A3E9" blockId="8.[151,1437,152,2034]" pageId="8" pageNumber="9">
We found
<taxonomicName id="00BAECF20D29755C6A06B65067D3A305" box="[321,433,1041,1066]" class="Insecta" family="Nymphalidae" genus="Dynastor" kingdom="Animalia" order="Lepidoptera" pageId="8" pageNumber="9" phylum="Arthropoda" rank="species" species="darius">
<emphasis id="F5CE4B630D29755C6A06B65067D3A305" box="[321,433,1041,1066]" italics="true" pageId="8" pageNumber="9">D. darius</emphasis>
</taxonomicName>
and
<taxonomicName id="00BAECF20D29755C6AABB650641DA305" box="[492,639,1041,1066]" class="Insecta" family="Nymphalidae" genus="Dynastor" kingdom="Animalia" order="Lepidoptera" pageId="8" pageNumber="9" phylum="Arthropoda" rank="species" species="napoleon">
<emphasis id="F5CE4B630D29755C6AABB650641DA305" box="[492,639,1041,1066]" italics="true" pageId="8" pageNumber="9">D. napoleon</emphasis>
</taxonomicName>
to be sister taxa, and two homoplasious transformations supported this clade: the presence of midleg tibial spurs (21:1), also present in
<taxonomicName id="00BAECF20D29755C68FBB679623CA37E" box="[956,1118,1080,1105]" class="Insecta" family="Nymphalidae" genus="Narope" kingdom="Animalia" order="Lepidoptera" pageId="8" pageNumber="9" phylum="Arthropoda" rank="species" species="panniculus">
<emphasis id="F5CE4B630D29755C68FBB679623CA37E" box="[956,1118,1080,1105]" italics="true" pageId="8" pageNumber="9">N. panniculus</emphasis>
</taxonomicName>
; and last segment of labial palpus approximately the same length as distal width of second segment (1:1,
<figureCitation id="5F818BF40D29755C6F14B61F62F9A357" box="[1107,1179,1118,1144]" captionStart="FIGURE 8" captionStartId="14.[151,255,1625,1649]" captionTargetBox="[157,1409,196,1582]" captionTargetId="figure@14.[151,1436,164,1735]" captionTargetPageId="14" captionText="FIGURE 8. Male characters. Scale bars, 1 mm. Labial palp in lateral view. (A) Bia actorion. (B) Brassolis sophorae. (C) Dynastor darius. Tegumen in dorsal view. (D) B. astyra. Genitalic capsule in lateral view. (E) D. darius. (F) B. sophorae. Tegumen and uncus in lateral view. (G) D. macrosiris. Phallus in lateral view. (H) D. napoleon. Juxta in ventral view. (I) B. astyra. (J) D. napoleon. Phallus in ventral view. (K) D. darius. (L) B. haenschi. (M) B. granadensis." httpUri="https://zenodo.org/record/188456/files/figure.png" pageId="8" pageNumber="9">Fig. 8</figureCitation>
C), present also in
<taxonomicName id="00BAECF20D29755C6EC5B61E669DA3B1" class="Insecta" family="Nymphalidae" genus="Brassolis" kingdom="Animalia" order="Lepidoptera" pageId="8" pageNumber="9" phylum="Arthropoda" rank="species" species="haenschi">
<emphasis id="F5CE4B630D29755C6EC5B61E669DA3B1" italics="true" pageId="8" pageNumber="9">B. haenschi</emphasis>
</taxonomicName>
. This clade had moderated values of Relative Bremer support and jackknife recuperation (50 and 52;
<figureCitation id="5F818BF40D29755C6BD0B6ED66BEA3E9" box="[151,220,1196,1222]" captionStart="FIGURE 4" captionStartId="10.[151,261,1680,1704]" captionTargetBox="[207,1385,425,1637]" captionTargetId="figure@10.[165,1425,391,1651]" captionTargetPageId="10" captionText="FIGURE 4. (A) Strict consensus tree from the analysis of 57 characters with K = 3. (B &amp; C) Two of the most parsimonious trees, the third tree had the same topology of the strict consensus. Relative Bremer support (on the left) and Jackknife values (on the right) are on the left hand of the branches." httpUri="https://zenodo.org/record/188452/files/figure.png" pageId="8" pageNumber="9">Fig. 4</figureCitation>
A).
</paragraph>
<paragraph id="C70597710D29755C6B82B69264EFA025" blockId="8.[151,1437,152,2034]" pageId="8" pageNumber="9">
The monophyly of
<taxonomicName id="00BAECF20D29755C6A8DB6926425A3C3" box="[458,583,1235,1260]" class="Insecta" family="Nymphalidae" genus="Brassolis" kingdom="Animalia" order="Lepidoptera" pageId="8" pageNumber="9" phylum="Arthropoda" rank="genus">
<emphasis id="F5CE4B630D29755C6A8DB6926425A3C3" box="[458,583,1235,1260]" italics="true" pageId="8" pageNumber="9">Brassolis</emphasis>
</taxonomicName>
was supported by 13 synapomorphies and three homoplasious transformations. The synapomorphies of
<taxonomicName id="00BAECF20D29755C69C5B6B86492A23D" box="[642,752,1273,1298]" class="Insecta" family="Nymphalidae" genus="Brassolis" kingdom="Animalia" order="Lepidoptera" pageId="8" pageNumber="9" phylum="Arthropoda" rank="genus">
<emphasis id="F5CE4B630D29755C69C5B6B86492A23D" box="[642,752,1273,1298]" italics="true" pageId="8" pageNumber="9">Brassolis</emphasis>
</taxonomicName>
were: absence of sensilla styloconica at the proboscis tip (2:0,
<figureCitation id="5F818BF40D29755C6B90B761677CA215" box="[215,286,1312,1338]" captionStart="FIGURE 3" captionStartId="6.[151,257,1913,1937]" captionTargetBox="[223,1316,798,1878]" captionTargetId="figure@6.[218,1348,783,1910]" captionTargetPageId="6" captionText="FIGURE 3. Brassolis species, in dorsal (left) and ventral (right) views (except for C). (A &amp; B) Male and female B. isthmia from Panama. (C) Male B. isthmia from Costa Rica. (D) Male of granadensis from Colombia. (E) Detail showing head and proboscis of B. isthmia from Panama. (F &amp; G) Male and female B. astyra from Rio de Janeiro (Brazil). (H) Male B. astyra from Rio Grande do Sul (Brazil)." httpUri="https://zenodo.org/record/188451/files/figure.png" pageId="8" pageNumber="9">Fig. 3</figureCitation>
E); in males, basal third of forewing costal margin slightly concave (3:1,
<figureCitation id="5F818BF40D29755C6FDFB7616282A215" box="[1176,1248,1312,1338]" captionStart="FIGURE 7" captionStartId="13.[151,255,1907,1931]" captionTargetBox="[202,1408,195,1834]" captionTargetId="figure@13.[151,1436,157,1880]" captionTargetPageId="13" captionText="FIGURE 7. Wing characters. (A) Brassolis dizini. (B) B. astyra. (C) B. haenschi. (D) Dynastor darius." httpUri="https://zenodo.org/record/188455/files/figure.png" pageId="8" pageNumber="9">Fig. 7</figureCitation>
A); male dorsal forewing postmedial band extended through discal cell (6:0,
<figureCitation id="5F818BF40D29755C6831B70765DCA24F" box="[886,958,1350,1376]" captionStart="FIGURE 7" captionStartId="13.[151,255,1907,1931]" captionTargetBox="[202,1408,195,1834]" captionTargetId="figure@13.[151,1436,157,1880]" captionTargetPageId="13" captionText="FIGURE 7. Wing characters. (A) Brassolis dizini. (B) B. astyra. (C) B. haenschi. (D) Dynastor darius." httpUri="https://zenodo.org/record/188455/files/figure.png" pageId="8" pageNumber="9">Fig. 7</figureCitation>
B); presence of a sent organ adjacent to male hindwing vein Cu2 (15:1,
<figureCitation id="5F818BF40D29755C694EB72C642FA2A8" box="[521,589,1389,1415]" captionStart="FIGURE 7" captionStartId="13.[151,255,1907,1931]" captionTargetBox="[202,1408,195,1834]" captionTargetId="figure@13.[151,1436,157,1880]" captionTargetPageId="13" captionText="FIGURE 7. Wing characters. (A) Brassolis dizini. (B) B. astyra. (C) B. haenschi. (D) Dynastor darius." httpUri="https://zenodo.org/record/188455/files/figure.png" pageId="8" pageNumber="9">Fig. 7</figureCitation>
A), in males, absence of a dorsal hindwing marginal line (18:0,
<figureCitation id="5F818BF40D29755C6E76B72C6317A2A8" box="[1329,1397,1389,1415]" captionStart="FIGURE 7" captionStartId="13.[151,255,1907,1931]" captionTargetBox="[202,1408,195,1834]" captionTargetId="figure@13.[151,1436,157,1880]" captionTargetPageId="13" captionText="FIGURE 7. Wing characters. (A) Brassolis dizini. (B) B. astyra. (C) B. haenschi. (D) Dynastor darius." httpUri="https://zenodo.org/record/188455/files/figure.png" pageId="8" pageNumber="9">Fig. 7</figureCitation>
A); in males, presence of dashed pattern of the distal portion of ventral hindwing discal cell (19:1,
<figureCitation id="5F818BF40D29755C6E6AB7D56314A281" box="[1325,1398,1428,1454]" captionStart="FIGURE 7" captionStartId="13.[151,255,1907,1931]" captionTargetBox="[202,1408,195,1834]" captionTargetId="figure@13.[151,1436,157,1880]" captionTargetPageId="13" captionText="FIGURE 7. Wing characters. (A) Brassolis dizini. (B) B. astyra. (C) B. haenschi. (D) Dynastor darius." httpUri="https://zenodo.org/record/188455/files/figure.png" pageId="8" pageNumber="9">Fig. 7</figureCitation>
C); presence of an anterolateral constriction of the tegumen (26:1,
<figureCitation id="5F818BF40D29755C6834B7FA65D5A2FA" box="[883,951,1467,1493]" captionStart="FIGURE 8" captionStartId="14.[151,255,1625,1649]" captionTargetBox="[157,1409,196,1582]" captionTargetId="figure@14.[151,1436,164,1735]" captionTargetPageId="14" captionText="FIGURE 8. Male characters. Scale bars, 1 mm. Labial palp in lateral view. (A) Bia actorion. (B) Brassolis sophorae. (C) Dynastor darius. Tegumen in dorsal view. (D) B. astyra. Genitalic capsule in lateral view. (E) D. darius. (F) B. sophorae. Tegumen and uncus in lateral view. (G) D. macrosiris. Phallus in lateral view. (H) D. napoleon. Juxta in ventral view. (I) B. astyra. (J) D. napoleon. Phallus in ventral view. (K) D. darius. (L) B. haenschi. (M) B. granadensis." httpUri="https://zenodo.org/record/188456/files/figure.png" pageId="8" pageNumber="9">Fig. 8</figureCitation>
D); internal surface of valva, basal region conspicuously less sclerotized than distal region (33:1); juxta not reaching the edge of the anellus (36:0,
<figureCitation id="5F818BF40D29755C6E2AB7A066C5A10D" captionStart="FIGURE 8" captionStartId="14.[151,255,1625,1649]" captionTargetBox="[157,1409,196,1582]" captionTargetId="figure@14.[151,1436,164,1735]" captionTargetPageId="14" captionText="FIGURE 8. Male characters. Scale bars, 1 mm. Labial palp in lateral view. (A) Bia actorion. (B) Brassolis sophorae. (C) Dynastor darius. Tegumen in dorsal view. (D) B. astyra. Genitalic capsule in lateral view. (E) D. darius. (F) B. sophorae. Tegumen and uncus in lateral view. (G) D. macrosiris. Phallus in lateral view. (H) D. napoleon. Juxta in ventral view. (I) B. astyra. (J) D. napoleon. Phallus in ventral view. (K) D. darius. (L) B. haenschi. (M) B. granadensis." httpUri="https://zenodo.org/record/188456/files/figure.png" pageId="8" pageNumber="9">
Fig.
<date id="B304B1B10D29755C6BD0B44966C5A10D" box="[151,167,1544,1570]" pageId="8" pageNumber="9">8</date>
</figureCitation>
I); presence of peg-like setae on semi-sclerotized lobe on left side of phallus (41:1,
<figureCitation id="5F818BF40D29755C6FE4B4496289A10D" box="[1187,1259,1544,1570]" captionStart="FIGURE 8" captionStartId="14.[151,255,1625,1649]" captionTargetBox="[157,1409,196,1582]" captionTargetId="figure@14.[151,1436,164,1735]" captionTargetPageId="14" captionText="FIGURE 8. Male characters. Scale bars, 1 mm. Labial palp in lateral view. (A) Bia actorion. (B) Brassolis sophorae. (C) Dynastor darius. Tegumen in dorsal view. (D) B. astyra. Genitalic capsule in lateral view. (E) D. darius. (F) B. sophorae. Tegumen and uncus in lateral view. (G) D. macrosiris. Phallus in lateral view. (H) D. napoleon. Juxta in ventral view. (I) B. astyra. (J) D. napoleon. Phallus in ventral view. (K) D. darius. (L) B. haenschi. (M) B. granadensis." httpUri="https://zenodo.org/record/188456/files/figure.png" pageId="8" pageNumber="9">Fig. 8</figureCitation>
K); absence of gnathos (49:0); fully sclerotized antrum (55:2); and posterior lobes of the papillae annales more developed ventrally (57:2,
<figureCitation id="5F818BF40D29755C6A18B41467CBA140" box="[351,425,1621,1647]" captionStart="FIGURE 9" captionStartId="15.[151,256,1456,1480]" captionTargetBox="[153,1412,201,1455]" captionTargetId="figure@15.[151,1436,127,1497]" captionTargetPageId="15" captionText="FIGURE 9. Female characters. Scale bars, 1 mm. Female distal tarsomere in ventral view (details of pulvillar process omitted). (A) Dynastor darius. Sterigma in ventral view. (B) D. darius. (C) B. isthmia. (D) B. astyra. (E) B. dinizi. Posterior lobes of papillae annales in lateral view. (F) Elymnias hypermnestra. (G) D. napoleon. (H) B. haenschi. Note that the lamella antevaginalis is not visible in C, D, and E due to the orientation of the drawing." httpUri="https://zenodo.org/record/188457/files/figure.png" pageId="8" pageNumber="9">Fig. 9</figureCitation>
H). The homoplasious transformations were: male midleg tarsus longer than tibia (22:0); presence of a semi-sclerotized lobe on the left side of phallus (40:1), which was also found in
<taxonomicName id="00BAECF20D29755C6E38B43D6680A193" class="Insecta" family="Nymphalidae" genus="Dynastor" kingdom="Animalia" order="Lepidoptera" pageId="8" pageNumber="9" phylum="Arthropoda" rank="species" species="darius">
<emphasis id="F5CE4B630D29755C6E38B43D6680A193" italics="true" pageId="8" pageNumber="9">D. darius</emphasis>
</taxonomicName>
and in one of the outgroup taxa (
<taxonomicName id="00BAECF20D29755C692BB4E265E0A193" box="[620,898,1699,1724]" class="Insecta" family="Nymphalidae" genus="Elymnias" kingdom="Animalia" order="Lepidoptera" pageId="8" pageNumber="9" phylum="Arthropoda" rank="species" species="hypermnestra">
<emphasis id="F5CE4B630D29755C692BB4E265E0A193" box="[620,898,1699,1724]" italics="true" pageId="8" pageNumber="9">Elymnias hypermnestra</emphasis>
</taxonomicName>
); and the transition between pedunculum and vinculum being straight (28:0,
<figureCitation id="5F818BF40D29755C695BB488643FA1CC" box="[540,605,1737,1763]" captionStart="FIGURE 8" captionStartId="14.[151,255,1625,1649]" captionTargetBox="[157,1409,196,1582]" captionTargetId="figure@14.[151,1436,164,1735]" captionTargetPageId="14" captionText="FIGURE 8. Male characters. Scale bars, 1 mm. Labial palp in lateral view. (A) Bia actorion. (B) Brassolis sophorae. (C) Dynastor darius. Tegumen in dorsal view. (D) B. astyra. Genitalic capsule in lateral view. (E) D. darius. (F) B. sophorae. Tegumen and uncus in lateral view. (G) D. macrosiris. Phallus in lateral view. (H) D. napoleon. Juxta in ventral view. (I) B. astyra. (J) D. napoleon. Phallus in ventral view. (K) D. darius. (L) B. haenschi. (M) B. granadensis." httpUri="https://zenodo.org/record/188456/files/figure.png" pageId="8" pageNumber="9">Fig 8</figureCitation>
F). Relative Bremer support and Jackknife recuperation values for
<taxonomicName id="00BAECF20D29755C6BD0B4B16766A026" box="[151,260,1776,1801]" class="Insecta" family="Nymphalidae" genus="Brassolis" kingdom="Animalia" order="Lepidoptera" pageId="8" pageNumber="9" phylum="Arthropoda" rank="genus">
<emphasis id="F5CE4B630D29755C6BD0B4B16766A026" box="[151,260,1776,1801]" italics="true" pageId="8" pageNumber="9">Brassolis</emphasis>
</taxonomicName>
were high (81 and 100;
<figureCitation id="5F818BF40D29755C6965B4B1640AA025" box="[546,616,1776,1802]" captionStart="FIGURE 4" captionStartId="10.[151,261,1680,1704]" captionTargetBox="[207,1385,425,1637]" captionTargetId="figure@10.[165,1425,391,1651]" captionTargetPageId="10" captionText="FIGURE 4. (A) Strict consensus tree from the analysis of 57 characters with K = 3. (B &amp; C) Two of the most parsimonious trees, the third tree had the same topology of the strict consensus. Relative Bremer support (on the left) and Jackknife values (on the right) are on the left hand of the branches." httpUri="https://zenodo.org/record/188452/files/figure.png" pageId="8" pageNumber="9">Fig. 4</figureCitation>
A).
</paragraph>
<paragraph id="C70597710D29755D6B82B5576581A79D" blockId="8.[151,1437,152,2034]" lastBlockId="9.[151,1437,152,990]" lastPageId="9" lastPageNumber="10" pageId="8" pageNumber="9">
Poor resolution of the strict consensus tree reflects the overall morphological similarity within
<taxonomicName id="00BAECF20D29755C6E69B55663FEA01F" box="[1326,1436,1815,1840]" class="Insecta" family="Nymphalidae" genus="Brassolis" kingdom="Animalia" order="Lepidoptera" pageId="8" pageNumber="9" phylum="Arthropoda" rank="genus">
<emphasis id="F5CE4B630D29755C6E69B55663FEA01F" box="[1326,1436,1815,1840]" italics="true" pageId="8" pageNumber="9">Brassolis</emphasis>
</taxonomicName>
(
<figureCitation id="5F818BF40D29755C6BE7B57C668AA078" box="[160,232,1853,1879]" captionStart="FIGURE 4" captionStartId="10.[151,261,1680,1704]" captionTargetBox="[207,1385,425,1637]" captionTargetId="figure@10.[165,1425,391,1651]" captionTargetPageId="10" captionText="FIGURE 4. (A) Strict consensus tree from the analysis of 57 characters with K = 3. (B &amp; C) Two of the most parsimonious trees, the third tree had the same topology of the strict consensus. Relative Bremer support (on the left) and Jackknife values (on the right) are on the left hand of the branches." httpUri="https://zenodo.org/record/188452/files/figure.png" pageId="8" pageNumber="9">Fig. 4</figureCitation>
A).
<emphasis id="F5CE4B630D29755C6A51B57C6498A079" box="[278,762,1853,1878]" italics="true" pageId="8" pageNumber="9">
<taxonomicName id="00BAECF20D29755C6A51B57C679DA079" box="[278,511,1853,1878]" class="Insecta" family="Nymphalidae" genus="Brassolis" kingdom="Animalia" order="Lepidoptera" pageId="8" pageNumber="9" phylum="Arthropoda" rank="species" species="sophorae">Brassolis sophorae</taxonomicName>
,
<taxonomicName id="00BAECF20D29755C694BB57C6410A079" box="[524,626,1853,1878]" class="Insecta" family="Nymphalidae" genus="Brassolis" kingdom="Animalia" order="Lepidoptera" pageId="8" pageNumber="9" phylum="Arthropoda" rank="species" species="dinizi">B. dinizi</taxonomicName>
,
<taxonomicName id="00BAECF20D29755C69C7B57C6498A079" box="[640,762,1853,1878]" class="Insecta" family="Nymphalidae" genus="Brassolis" kingdom="Animalia" order="Lepidoptera" pageId="8" pageNumber="9" phylum="Arthropoda" rank="species" species="isthmia">B. isthmia</taxonomicName>
</emphasis>
and
<taxonomicName id="00BAECF20D29755C6872B57C65A2A079" box="[821,960,1853,1878]" class="Insecta" family="Nymphalidae" genus="Brassolis" kingdom="Animalia" order="Lepidoptera" pageId="8" pageNumber="9" phylum="Arthropoda" rank="species" species="haenschi">
<emphasis id="F5CE4B630D29755C6872B57C65A2A079" box="[821,960,1853,1878]" italics="true" pageId="8" pageNumber="9">B. haenschi</emphasis>
</taxonomicName>
formed an unresolved clade (sophoraeclade) supported by one homoplasious transformation: presence of a dorsal hindwing postmedial band in males (17:1,
<figureCitation id="5F818BF40D29755C6A77B5CA6717A08A" box="[304,373,1931,1957]" captionStart="FIGURE 7" captionStartId="13.[151,255,1907,1931]" captionTargetBox="[202,1408,195,1834]" captionTargetId="figure@13.[151,1436,157,1880]" captionTargetPageId="13" captionText="FIGURE 7. Wing characters. (A) Brassolis dizini. (B) B. astyra. (C) B. haenschi. (D) Dynastor darius." httpUri="https://zenodo.org/record/188455/files/figure.png" pageId="8" pageNumber="9">Fig. 7</figureCitation>
A). Alternative groupings of these species were represented in the other two trees (
<figureCitation id="5F818BF40D29755C6E05B5CA63EAA08A" box="[1346,1416,1931,1957]" captionStart="FIGURE 4" captionStartId="10.[151,261,1680,1704]" captionTargetBox="[207,1385,425,1637]" captionTargetId="figure@10.[165,1425,391,1651]" captionTargetPageId="10" captionText="FIGURE 4. (A) Strict consensus tree from the analysis of 57 characters with K = 3. (B &amp; C) Two of the most parsimonious trees, the third tree had the same topology of the strict consensus. Relative Bremer support (on the left) and Jackknife values (on the right) are on the left hand of the branches." httpUri="https://zenodo.org/record/188452/files/figure.png" pageId="8" pageNumber="9">Fig. 4</figureCitation>
B and C). In all resolutions, however,
<taxonomicName id="00BAECF20D29755C691EB5F06571A0E5" box="[601,787,1969,1994]" class="Insecta" family="Nymphalidae" genus="Brassolis" kingdom="Animalia" order="Lepidoptera" pageId="8" pageNumber="9" phylum="Arthropoda" rank="species" species="granadensis">
<emphasis id="F5CE4B630D29755C691EB5F06571A0E5" box="[601,787,1969,1994]" italics="true" pageId="8" pageNumber="9">B. granadensis</emphasis>
</taxonomicName>
was the sister taxon of this clade, supported by one homoplasious transformation: the presence of long setae on a semi-sclerotized lobe on the left side of phallus (43:1,
<figureCitation id="5F818BF40D28755D6BA4B2D96745A79D" box="[227,295,152,178]" captionStart="FIGURE 8" captionStartId="14.[151,255,1625,1649]" captionTargetBox="[157,1409,196,1582]" captionTargetId="figure@14.[151,1436,164,1735]" captionTargetPageId="14" captionText="FIGURE 8. Male characters. Scale bars, 1 mm. Labial palp in lateral view. (A) Bia actorion. (B) Brassolis sophorae. (C) Dynastor darius. Tegumen in dorsal view. (D) B. astyra. Genitalic capsule in lateral view. (E) D. darius. (F) B. sophorae. Tegumen and uncus in lateral view. (G) D. macrosiris. Phallus in lateral view. (H) D. napoleon. Juxta in ventral view. (I) B. astyra. (J) D. napoleon. Phallus in ventral view. (K) D. darius. (L) B. haenschi. (M) B. granadensis." httpUri="https://zenodo.org/record/188456/files/figure.png" pageId="9" pageNumber="10">Fig. 8</figureCitation>
L). Finally,
<taxonomicName id="00BAECF20D28755D6AF5B2D96479A79E" box="[434,539,152,177]" class="Insecta" family="Nymphalidae" genus="Brassolis" kingdom="Animalia" order="Lepidoptera" pageId="9" pageNumber="10" phylum="Arthropoda" rank="species" species="astyra">
<emphasis id="F5CE4B630D28755D6AF5B2D96479A79E" box="[434,539,152,177]" italics="true" pageId="9" pageNumber="10">B. astyra</emphasis>
</taxonomicName>
appears as sister to all other
<taxonomicName id="00BAECF20D28755D6828B2D965BEA79E" box="[879,988,152,177]" class="Insecta" family="Nymphalidae" genus="Brassolis" kingdom="Animalia" order="Lepidoptera" pageId="9" pageNumber="10" phylum="Arthropoda" rank="genus">
<emphasis id="F5CE4B630D28755D6828B2D965BEA79E" box="[879,988,152,177]" italics="true" pageId="9" pageNumber="10">Brassolis</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="C70597710D28755D6B82B2FF674CA6EF" blockId="9.[151,1437,152,990]" pageId="9" pageNumber="10">
To identify characters that produced conflicting topologies within the sophorae-clade, we did a search without collapsing zero-length branches, and optimized both ambiguous and unambiguous transformations onto the obtained trees. The results were five alternative topologies for the sophorae-clade (
<figureCitation id="5F818BF40D28755D6F9AB34D6345A609" box="[1245,1319,268,294]" captionStart="FIGURE 5" captionStartId="11.[151,261,1839,1863]" captionTargetBox="[192,1377,188,1825]" captionTargetId="figure@11.[151,1436,125,1893]" captionTargetPageId="11" captionText="FIGURE 5. (A E) Five resolutions obtained from a search without collapsing zero-length branches showing optimization of both unambiguous and ambiguous transformations. Symbols follow the legend in Fig. 4. The horizontal bars above the trees indicate the sophorae-clade." httpUri="https://zenodo.org/record/188453/files/figure.png" pageId="9" pageNumber="10">Fig. 5</figureCitation>
). In these five resolved sub-trees all characters producing ambiguous optimizations (11, 12, 14, 16, 20, 25, 54) but one (54) were coloration characters, and all but one (12) were polymorphic. Trees in
<figureCitation id="5F818BF40D28755D6F1FB31862FCA65C" box="[1112,1182,345,371]" captionStart="FIGURE 5" captionStartId="11.[151,261,1839,1863]" captionTargetBox="[192,1377,188,1825]" captionTargetId="figure@11.[151,1436,125,1893]" captionTargetPageId="11" captionText="FIGURE 5. (A E) Five resolutions obtained from a search without collapsing zero-length branches showing optimization of both unambiguous and ambiguous transformations. Symbols follow the legend in Fig. 4. The horizontal bars above the trees indicate the sophorae-clade." httpUri="https://zenodo.org/record/188453/files/figure.png" pageId="9" pageNumber="10">Fig. 5</figureCitation>
show
<taxonomicName id="00BAECF20D28755D6FABB3186319A65D" box="[1260,1403,345,370]" class="Insecta" family="Nymphalidae" genus="Brassolis" kingdom="Animalia" order="Lepidoptera" pageId="9" pageNumber="10" phylum="Arthropoda" rank="species" species="sophorae">
<emphasis id="F5CE4B630D28755D6FABB3186319A65D" box="[1260,1403,345,370]" italics="true" pageId="9" pageNumber="10">B. sophorae</emphasis>
</taxonomicName>
as sister to each of the species in the sophorae-clade (
<figureCitation id="5F818BF40D28755D69B7B3C16557A6B5" box="[752,821,384,410]" captionStart="FIGURE 5" captionStartId="11.[151,261,1839,1863]" captionTargetBox="[192,1377,188,1825]" captionTargetId="figure@11.[151,1436,125,1893]" captionTargetPageId="11" captionText="FIGURE 5. (A E) Five resolutions obtained from a search without collapsing zero-length branches showing optimization of both unambiguous and ambiguous transformations. Symbols follow the legend in Fig. 4. The horizontal bars above the trees indicate the sophorae-clade." httpUri="https://zenodo.org/record/188453/files/figure.png" pageId="9" pageNumber="10">Fig. 5</figureCitation>
CE) or as sister to species groups within this clade (
<figureCitation id="5F818BF40D28755D6BD8B3E76686A6EF" box="[159,228,422,448]" captionStart="FIGURE 5" captionStartId="11.[151,261,1839,1863]" captionTargetBox="[192,1377,188,1825]" captionTargetId="figure@11.[151,1436,125,1893]" captionTargetPageId="11" captionText="FIGURE 5. (A E) Five resolutions obtained from a search without collapsing zero-length branches showing optimization of both unambiguous and ambiguous transformations. Symbols follow the legend in Fig. 4. The horizontal bars above the trees indicate the sophorae-clade." httpUri="https://zenodo.org/record/188453/files/figure.png" pageId="9" pageNumber="10">Fig. 5</figureCitation>
AB).
</paragraph>
<paragraph id="C70597710D28755D6B82B38C6270A587" blockId="9.[151,1437,152,990]" pageId="9" pageNumber="10">
Given that polymorphic characters supported ambiguous optimizations within the sophorae-clade, an analysis was done excluding nine polymorphic characters (Appendix 3). Characters 11, 13, 14, 16, 20, 25, 43 and 54 were polymorphic for
<taxonomicName id="00BAECF20D28755D6ABDB05B64E8A51C" box="[506,650,538,563]" class="Insecta" family="Nymphalidae" genus="Brassolis" kingdom="Animalia" order="Lepidoptera" pageId="9" pageNumber="10" phylum="Arthropoda" rank="species" species="sophorae">
<emphasis id="F5CE4B630D28755D6ABDB05B64E8A51C" box="[506,650,538,563]" italics="true" pageId="9" pageNumber="10">B. sophorae</emphasis>
</taxonomicName>
, and character 17 was polymorphic for
<taxonomicName id="00BAECF20D28755D6F22B05B62BCA51C" box="[1125,1246,538,563]" class="Insecta" family="Nymphalidae" genus="Brassolis" kingdom="Animalia" order="Lepidoptera" pageId="9" pageNumber="10" phylum="Arthropoda" rank="species" species="isthmia">
<emphasis id="F5CE4B630D28755D6F22B05B62BCA51C" box="[1125,1246,538,563]" italics="true" pageId="9" pageNumber="10">B. isthmia</emphasis>
</taxonomicName>
, and also for
<taxonomicName id="00BAECF20D28755D6EC4B05B6680A575" class="Insecta" family="Nymphalidae" genus="Brassolis" kingdom="Animalia" order="Lepidoptera" pageId="9" pageNumber="10" phylum="Arthropoda" rank="species" species="astyra">
<emphasis id="F5CE4B630D28755D6EC4B05B6680A575" italics="true" pageId="9" pageNumber="10">B. astyra</emphasis>
</taxonomicName>
(outside the sophorae-clade). Two trees were found that differed greatly in resolution, and the strict consensus of these trees showed the sophorae-clade as a polytomy. We therefore concluded that the exclusion of polymorphic characters did not improve our results (trees not illustrated).
</paragraph>
<paragraph id="C70597710D28755D6B82B0F46694A4F1" blockId="9.[151,1437,152,990]" pageId="9" pageNumber="10">
As noted above,
<taxonomicName id="00BAECF20D28755D6AD1B0F46449A5E1" box="[406,555,693,718]" class="Insecta" family="Nymphalidae" genus="Brassolis" kingdom="Animalia" order="Lepidoptera" pageId="9" pageNumber="10" phylum="Arthropoda" rank="species" species="sophorae">
<emphasis id="F5CE4B630D28755D6AD1B0F46449A5E1" box="[406,555,693,718]" italics="true" pageId="9" pageNumber="10">B. sophorae</emphasis>
</taxonomicName>
was polymorphic for eight characters, and we therefore asked whether splitting
<taxonomicName id="00BAECF20D28755D6A47B09D67F2A5DA" box="[256,400,732,757]" class="Insecta" family="Nymphalidae" genus="Brassolis" kingdom="Animalia" order="Lepidoptera" pageId="9" pageNumber="10" phylum="Arthropoda" rank="species" species="sophorae">
<emphasis id="F5CE4B630D28755D6A47B09D67F2A5DA" box="[256,400,732,757]" italics="true" pageId="9" pageNumber="10">B. sophorae</emphasis>
</taxonomicName>
into monomorphic terminals would improve tree resolution and would provide a more reliable assessment of relationships. To investigate this, we used male-female pairs (whenever possible) collected in eight countries as terminals. This sampling attempted to represent the
<typeStatus id="180129D30D28755D6F37B16862C1A46C" box="[1136,1187,809,835]" pageId="9" pageNumber="10">type</typeStatus>
locality (country) of most valid subspecies listed by
<bibRefCitation id="A32BEA800D28755D694AB111648DA445" author="Casagrande" box="[525,751,848,874]" pageId="9" pageNumber="17" refString="Casagrande, M. M. (2004) Brassolini. Atlas of Neotropical Lepidoptera (ed. By J. B. Heppner), Part 4 A Checklist (ed. By G. Lamas), pp. 201 - 205. Association of Tropical Lepidoptera / Scientific Publishers, Gainesville, FL." type="book chapter" year="2004">Casagrande (2004)</bibRefCitation>
, therefore approximating a range of character variation of
<taxonomicName id="00BAECF20D28755D6BD0B1366748A4BF" box="[151,298,887,912]" class="Insecta" family="Nymphalidae" genus="Brassolis" kingdom="Animalia" order="Lepidoptera" pageId="9" pageNumber="10" phylum="Arthropoda" rank="species" species="sophorae">
<emphasis id="F5CE4B630D28755D6BD0B1366748A4BF" box="[151,298,887,912]" italics="true" pageId="9" pageNumber="10">B. sophorae</emphasis>
</taxonomicName>
. Separating
<taxonomicName id="00BAECF20D28755D6A83B136643AA4BF" box="[452,600,887,912]" class="Insecta" family="Nymphalidae" genus="Brassolis" kingdom="Animalia" order="Lepidoptera" pageId="9" pageNumber="10" phylum="Arthropoda" rank="species" species="sophorae">
<emphasis id="F5CE4B630D28755D6A83B136643AA4BF" box="[452,600,887,912]" italics="true" pageId="9" pageNumber="10">B. sophorae</emphasis>
</taxonomicName>
into eight terminals (putative subspecies) showed that all species of
<taxonomicName id="00BAECF20D28755D6BD0B1DC6766A499" box="[151,260,925,950]" class="Insecta" family="Nymphalidae" genus="Brassolis" kingdom="Animalia" order="Lepidoptera" pageId="9" pageNumber="10" phylum="Arthropoda" rank="genus">
<emphasis id="F5CE4B630D28755D6BD0B1DC6766A499" box="[151,260,925,950]" italics="true" pageId="9" pageNumber="10">Brassolis</emphasis>
</taxonomicName>
are associated with one or more subspecies of
<taxonomicName id="00BAECF20D28755D6877B1DC65FFA499" box="[816,925,925,950]" class="Insecta" family="Nymphalidae" genus="Brassolis" kingdom="Animalia" order="Lepidoptera" pageId="9" pageNumber="10" phylum="Arthropoda" rank="species" species="sophorae">
<emphasis id="F5CE4B630D28755D6877B1DC65FFA499" box="[816,925,925,950]" italics="true" pageId="9" pageNumber="10">sophorae</emphasis>
</taxonomicName>
; i.e., the sophorae-clade was not recovered (
<figureCitation id="5F818BF40D28755D6BD8B1856685A4F1" box="[159,231,964,990]" captionStart="FIGURE 6" captionStartId="12.[151,255,1117,1141]" captionTargetBox="[251,1301,191,1116]" captionTargetId="figure@12.[251,1335,137,1143]" captionTargetPageId="12" captionText="FIGURE 6. Strict consensus of three trees found when B. sophorae and B. astyra subspecies were used as terminals." httpUri="https://zenodo.org/record/188454/files/figure.png" pageId="9" pageNumber="10">Fig. 6</figureCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>