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<mods:title id="8A71D4CD831A2FE2D1E5C45B2FE149A0">A snow-dwelling tropical buưerfly? An unprecedented discovery of a new genus of the Pedaliodes clade in an extreme, high-altitude Andean environment (Lepidoptera: Nymphalidae: Satyrinae)</mods:title>
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<emphasis id="ADEFDE13FFF7FFF1FF2AD245B1688200" bold="true" box="[228,346,1840,1867]" italics="true" pageId="3" pageNumber="4">Nivaliodes</emphasis>
Pyrcz &amp; Espeland
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<emphasis id="ADEFDE13FFF7FFF1FFBFD223B0ED8225" bold="true" box="[113,223,1878,1902]" italics="true" pageId="3" pageNumber="4">Nivaliodes</emphasis>
Pyrcz &amp; Espeland
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species:
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<emphasis id="ADEFDE13FFF7FFF1FF25D2DAB18F828C" box="[235,445,1967,1991]" italics="true" pageId="3" pageNumber="4">Nivaliodes negrobueno</emphasis>
Pyrcz, Fåhraeus &amp; Boyer
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<emphasis id="ADEFDE13FFF7FFF1FC39D3E0B4D483E5" box="[1015,1254,1684,1710]" italics="true" pageId="3" pageNumber="4">Diagnosis and description</emphasis>
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Adults of medium size, with a FW with costa length between 21 and
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, with brown or blackish brown FWD and HWD, except for a white postdiscal costal patch in the FW, and with an almost uniform underside with some faint white scaling, blackbrown FW and black HW freely sprinkled by snow-white scales, HWV median costal patch whitish. No androconial patch, except for a few vestigial androconial scales detected under the scanning electron microscope in one of the three known species,
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<emphasis id="ADEFDE13FFF7FFF1FC92D2C0B3CE8286" box="[860,1020,1973,1997]" italics="true" pageId="3" pageNumber="4">Nivaliodes puriq</emphasis>
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(
<figureCitation id="07A01E84FFF7FFF1FBC2D2C0B47F8286" box="[1036,1101,1973,1997]" captionStart="Figure 7" captionStartId="11.[113,178,1772,1796]" captionTargetBox="[260,1313,149,1737]" captionTargetId="figure-7@11.[253,1320,144,1744]" captionTargetPageId="11" captionText="Figure 7. Scanning electron microscopy images of Nivaliodes negrobueno scales.A, scale-covered wing membrane, with visible discal veins also covered by ground scales. B, elongated cover scales with hairy scales. C, two morphotypes of scales: elongated cover scales and wide, shovel-shaped ground scales. D, elongated cover scales and hairy scales. E, elongated cover scale with pointed apical termini.F, shovel- like cover scale with undulating terminus. G, detail of cover scale micro- and structures, median area. H, detail of cover scale micro- and nanostructures, apical area." pageId="3" pageNumber="4">Figs. 7</figureCitation>
,
<figureCitation id="07A01E84FFF7FFF1FB96D2C0B4578286" box="[1112,1125,1973,1997]" captionStart="Figure 8" captionStartId="12.[129,194,1771,1795]" captionTargetBox="[275,1328,148,1736]" captionTargetId="figure-9@12.[268,1335,143,1743]" captionTargetPageId="12" captionText="Figure 8. Scanning electron microscopy images of Nivaliodes puriq scales.A, scale-covered wing membrane with visible veins free of scales.B, transformed cover scales with sparse hair-like androconial scales.C, transformed cover scales with elongated shape on top, shovel-like ground scales below.D, transformed cover scales and sparse androconial scales. E, transformed cover scale with pointed termini.F, fragments of a transformed cover scale and an androconial scale. G, detail of terminal part of a hair-like androconial scale. H, micro- and nanostructures in the transformed cover scale, median area." pageId="3" pageNumber="4">8</figureCitation>
). Wing venation similar to other genera of the
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<emphasis id="ADEFDE13FFF0FFF6FED4D3BCB1B283AA" box="[282,384,1737,1761]" italics="true" pageId="4" pageNumber="5">Pedaliodes</emphasis>
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clade (see:
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Pyrcz
<emphasis id="ADEFDE13FFF0FFF6FD83D3BFB2B383AA" box="[589,641,1737,1761]" italics="true" pageId="4" pageNumber="5">et al.</emphasis>
2023
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). Eyes densely hairy; palpi two times the length of head; antennae reaching two-fisshs the length of costa, slender, mostly naked, with a club formed gradually, composed of 11 flagellomeres, flattened dorsoventrally.
</paragraph>
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<emphasis id="ADEFDE13FFF0FFF6FF4FD3F4B0EB83D2" bold="true" box="[129,217,1665,1689]" pageId="4" pageNumber="5">Figure 1.</emphasis>
Maximum likelihood tree of
<taxonomicName id="589B7982FFF0FFF6FE22D3F4B26083D2" authorityName="Pyrcz &amp; Mahecha-J. &amp; Boyer &amp; Lachowska-Cielik &amp; Cerdeña &amp; Farfán &amp; Garlacz &amp; Lorenc-Brudecka &amp; Bálint &amp; Fåhraeus &amp; Zając-Garlacz &amp; Espeland" authorityYear="2024" box="[492,594,1665,1689]" class="Insecta" family="Boreidae" genus="Nivaliodes" kingdom="Animalia" order="Mecoptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="genus">
<emphasis id="ADEFDE13FFF0FFF6FE22D3F4B26083D2" bold="true" box="[492,594,1665,1689]" italics="true" pageId="4" pageNumber="5">Nivaliodes</emphasis>
</taxonomicName>
and other
<taxonomicName id="589B7982FFF0FFF6FD79D3F4B30683D2" box="[695,820,1665,1689]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="subTribe" subTribe="Pronophilina">Pronophilina</taxonomicName>
inferred in IQ-TREE based on target enrichment data and
<emphasis id="ADEFDE13FFF0FFF6FA90D3F7B5B583D2" box="[1374,1415,1666,1689]" italics="true" pageId="4" pageNumber="5">COI</emphasis>
.
</paragraph>
</caption>
<paragraph id="9F240201FFF0FFF6FF4FD2F5B4538215" blockId="4.[129,778,1920,1976]" lastBlockId="4.[825,1474,1737,1886]" pageId="4" pageNumber="5">
<emphasis id="ADEFDE13FFF0FFF6FF4FD2F5B12B82D3" box="[129,281,1920,1944]" italics="true" pageId="4" pageNumber="5">Male genitalia:</emphasis>
With stout uncus and subunci and, contrary to the species of
<taxonomicName id="589B7982FFF0FFF6FEF4D2EAB19282FC" box="[314,416,1951,1975]" class="Insecta" family="Nymphalidae" genus="Pedaliodes" kingdom="Animalia" order="Lepidoptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="genus">
<emphasis id="ADEFDE13FFF0FFF6FEF4D2EAB19282FC" box="[314,416,1951,1975]" italics="true" pageId="4" pageNumber="5">Pedaliodes</emphasis>
</taxonomicName>
, the valva is devoid of any dorsal process, instead with a humped, either slightly irregular or serrate dorsum, and with a short, straight, and wide, tubular aedeagus, differing from the contorted and usually considerably longer aedeagus of
<taxonomicName id="589B7982FFF0FFF6FC32D252B4508274" box="[1020,1122,1831,1855]" class="Insecta" family="Nymphalidae" genus="Pedaliodes" kingdom="Animalia" order="Lepidoptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="genus">
<emphasis id="ADEFDE13FFF0FFF6FC32D252B4508274" box="[1020,1122,1831,1855]" italics="true" pageId="4" pageNumber="5">Pedaliodes</emphasis>
</taxonomicName>
, bearing a sharp apical ending and a widened proximal opening.
</paragraph>
<paragraph id="9F240201FFF0FFF7FCF7D2F5B08A8311" blockId="4.[825,1475,1920,1975]" lastBlockId="5.[113,763,1477,1626]" lastPageId="5" lastPageNumber="6" pageId="4" pageNumber="5">
<emphasis id="ADEFDE13FFF0FFF6FCF7D2F5B3EC82D3" box="[825,990,1920,1944]" italics="true" pageId="4" pageNumber="5">Female genitalia:</emphasis>
Papillae anales prominent, postvaginal lamella wide and strongly sclerotized with a smooth surface, enclosing a spacious antrum; ductus bursae mildly sclerotized, medium long and very wide, opening gradually into a large, pear-like corpus bursae, with a smooth surface, no signa, unlike the species of
<taxonomicName id="589B7982FFF1FFF7FFBFD356B0E18370" box="[113,211,1571,1595]" class="Insecta" family="Nymphalidae" genus="Pedaliodes" kingdom="Animalia" order="Lepidoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="genus">
<emphasis id="ADEFDE13FFF1FFF7FFBFD356B0E18370" box="[113,211,1571,1595]" italics="true" pageId="5" pageNumber="6">Pedaliodes</emphasis>
</taxonomicName>
, which invariably have two parallel signa on corpus bursae.
</paragraph>
<caption id="CBE45289FFF1FFF7FFBFD012B2E180CB" box="[113,723,1383,1408]" pageId="5" pageNumber="6" startId="5.[113,178,1383,1407]" targetBox="[149,1419,146,1351]" targetPageId="5" targetType="figure">
<paragraph id="9F240201FFF1FFF7FFBFD012B2E180CB" blockId="5.[113,723,1383,1408]" box="[113,723,1383,1408]" pageId="5" pageNumber="6">
<emphasis id="ADEFDE13FFF1FFF7FFBFD012B0FB80CB" bold="true" box="[113,201,1383,1408]" pageId="5" pageNumber="6">Figure 2.</emphasis>
Divergence analysis of
<taxonomicName id="589B7982FFF1FFF7FE6AD012B2388034" authorityName="Pyrcz &amp; Mahecha-J. &amp; Boyer &amp; Lachowska-Cielik &amp; Cerdeña &amp; Farfán &amp; Garlacz &amp; Lorenc-Brudecka &amp; Bálint &amp; Fåhraeus &amp; Zając-Garlacz &amp; Espeland" authorityYear="2024" box="[420,522,1383,1407]" class="Insecta" family="Boreidae" genus="Nivaliodes" kingdom="Animalia" order="Mecoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="genus">
<emphasis id="ADEFDE13FFF1FFF7FE6AD012B2388034" bold="true" box="[420,522,1383,1407]" italics="true" pageId="5" pageNumber="6">Nivaliodes</emphasis>
</taxonomicName>
inferred in BEAST2.
</paragraph>
</caption>
<paragraph id="9F240201FFF1FFF7FEACD3F5B23B83D1" blockId="5.[113,763,1664,1977]" box="[354,521,1664,1690]" pageId="5" pageNumber="6">
<emphasis id="ADEFDE13FFF1FFF7FEACD3F5B23B83D1" box="[354,521,1664,1690]" italics="true" pageId="5" pageNumber="6">Scale morphology</emphasis>
</paragraph>
<paragraph id="9F240201FFF1FFF7FFBFD3D3B3D180B7" blockId="5.[113,763,1664,1977]" lastBlockId="5.[810,1459,1477,1532]" pageId="5" pageNumber="6">
Uppersides of wings are overlaid with two thickly packed layers of flat, cover scales, the top layer with predominantly elongated scales with dentate apices, and the bottom layer with scarcer, wide scales with gently undulating apices, visible only when top layer scales are removed; long, hairy scales are rather sparse and concentrate along outer areas of wings, fringes composed of elongated brushy scales. There are no specialized androconial scales. Scale ultramorphology is homogeneous, with the body of the individual scales being structured by longitudinal ridges and cross ribs connected by trabeculae, with no apparent lower-level lamina (
<figureCitation id="07A01E84FFF1FFF7FCB1D091B38980B7" box="[895,955,1508,1532]" captionStart="Figure 7" captionStartId="11.[113,178,1772,1796]" captionTargetBox="[260,1313,149,1737]" captionTargetId="figure-7@11.[253,1320,144,1744]" captionTargetPageId="11" captionText="Figure 7. Scanning electron microscopy images of Nivaliodes negrobueno scales.A, scale-covered wing membrane, with visible discal veins also covered by ground scales. B, elongated cover scales with hairy scales. C, two morphotypes of scales: elongated cover scales and wide, shovel-shaped ground scales. D, elongated cover scales and hairy scales. E, elongated cover scale with pointed apical termini.F, shovel- like cover scale with undulating terminus. G, detail of cover scale micro- and structures, median area. H, detail of cover scale micro- and nanostructures, apical area." pageId="5" pageNumber="6">Figs 7</figureCitation>
,
<figureCitation id="07A01E84FFF1FFF7FC08D091B3E180B7" box="[966,979,1508,1532]" captionStart="Figure 8" captionStartId="12.[129,194,1771,1795]" captionTargetBox="[275,1328,148,1736]" captionTargetId="figure-9@12.[268,1335,143,1743]" captionTargetPageId="12" captionText="Figure 8. Scanning electron microscopy images of Nivaliodes puriq scales.A, scale-covered wing membrane with visible veins free of scales.B, transformed cover scales with sparse hair-like androconial scales.C, transformed cover scales with elongated shape on top, shovel-like ground scales below.D, transformed cover scales and sparse androconial scales. E, transformed cover scale with pointed termini.F, fragments of a transformed cover scale and an androconial scale. G, detail of terminal part of a hair-like androconial scale. H, micro- and nanostructures in the transformed cover scale, median area." pageId="5" pageNumber="6">8</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="D781518AFFF1FFF7FBF2D369B3AA83D2" pageId="5" pageNumber="6" type="etymology">
<paragraph id="9F240201FFF1FFF7FBF2D369B490837D" blockId="5.[810,1459,1564,1689]" box="[1084,1186,1564,1590]" pageId="5" pageNumber="6">
<emphasis id="ADEFDE13FFF1FFF7FBF2D369B490837D" box="[1084,1186,1564,1590]" italics="true" pageId="5" pageNumber="6">Etymology</emphasis>
</paragraph>
<paragraph id="9F240201FFF1FFF7FCE4D337B3AA83D2" blockId="5.[810,1459,1564,1689]" pageId="5" pageNumber="6">
A compound noun:
<emphasis id="ADEFDE13FFF1FFF7FBC6D337B4778311" box="[1032,1093,1602,1626]" italics="true" pageId="5" pageNumber="6">Nival-</emphasis>
, root of
<emphasis id="ADEFDE13FFF1FFF7FB66D336B4D68311" box="[1192,1252,1603,1626]" italics="true" pageId="5" pageNumber="6">snowy</emphasis>
in Latin, -
<emphasis id="ADEFDE13FFF1FFF7FA98D337B5BE8311" box="[1366,1420,1602,1626]" italics="true" pageId="5" pageNumber="6">liodes</emphasis>
, an allusion to
<taxonomicName id="589B7982FFF1FFF7FC51D314B4378332" box="[927,1029,1633,1657]" class="Insecta" family="Nymphalidae" genus="Pedaliodes" kingdom="Animalia" order="Lepidoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="genus">
<emphasis id="ADEFDE13FFF1FFF7FC51D314B4378332" box="[927,1029,1633,1657]" italics="true" pageId="5" pageNumber="6">Pedaliodes</emphasis>
</taxonomicName>
. Noun grammatical gender is considered masculine.
</paragraph>
</subSubSection>
<subSubSection id="D781518AFFF1FFF5FC32D3CDB297840E" lastPageId="7" lastPageNumber="8" pageId="5" pageNumber="6" type="discussion">
<paragraph id="9F240201FFF1FFF7FC32D3CDB4D38399" blockId="5.[1020,1249,1720,1746]" box="[1020,1249,1720,1746]" pageId="5" pageNumber="6">
<emphasis id="ADEFDE13FFF1FFF7FC32D3CDB4D38399" box="[1020,1249,1720,1746]" italics="true" pageId="5" pageNumber="6">Phylogeny and radiation</emphasis>
</paragraph>
<paragraph id="9F240201FFF1FFF4FCE4D3AAB39E8287" blockId="5.[810,1460,1759,1971]" lastBlockId="6.[825,1474,1878,1996]" lastPageId="6" lastPageNumber="7" pageId="5" pageNumber="6">
The genus
<taxonomicName id="589B7982FFF1FFF7FC65D3AAB42383BC" authorityName="Pyrcz &amp; Mahecha-J. &amp; Boyer &amp; Lachowska-Cielik &amp; Cerdeña &amp; Farfán &amp; Garlacz &amp; Lorenc-Brudecka &amp; Bálint &amp; Fåhraeus &amp; Zając-Garlacz &amp; Espeland" authorityYear="2024" box="[939,1041,1759,1783]" class="Insecta" family="Boreidae" genus="Nivaliodes" kingdom="Animalia" order="Mecoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="genus">
<emphasis id="ADEFDE13FFF1FFF7FC65D3AAB42383BC" box="[939,1041,1759,1783]" italics="true" pageId="5" pageNumber="6">Nivaliodes</emphasis>
</taxonomicName>
is fully supported and sister to the genus
<taxonomicName id="589B7982FFF1FFF7FCBED38BB496825D" authority="Forster, 1964" authorityName="Forster" authorityYear="1964" box="[880,1188,1790,1814]" class="Insecta" family="Nymphalidae" genus="Pherepedaliodes" kingdom="Animalia" order="Lepidoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="genus">
<emphasis id="ADEFDE13FFF1FFF7FCBED38BB43E825D" box="[880,1036,1790,1814]" italics="true" pageId="5" pageNumber="6">Pherepedaliodes</emphasis>
<bibRefCitation id="FB0A7FF0FFF1FFF7FBD9D38AB496825D" author="Forster W &amp; Lepidoptera III" box="[1047,1188,1790,1814]" pageId="5" pageNumber="6" pagination="51 - 188" refId="ref11726" refString="Forster W. Beitrage zur Kenntnis der Insektenfauna Boliviens, XIX. Lepidoptera III. Satyridae, Veroffentlichungen Zoologischen Staatssammlung Munchen 1964; 8: 51 - 188." type="book chapter" year="1964">Forster, 1964</bibRefCitation>
</taxonomicName>
(
<typeStatus id="4020BCA3FFF1FFF7FB78D38BB4D5825D" box="[1206,1255,1790,1814]" pageId="5" pageNumber="6">type</typeStatus>
species:
<taxonomicName id="589B7982FFF1FFF7FA83D38BB4F0827D" authority="Grose-Smith &amp; Kirby, 1894" authorityName="Grose-Smith &amp; Kirby" authorityYear="1894" class="Insecta" family="Nymphalidae" genus="Pedaliodes" kingdom="Animalia" order="Lepidoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="species" species="pheretiades">
<emphasis id="ADEFDE13FFF1FFF7FA83D38BB3A5827E" italics="true" pageId="5" pageNumber="6">Pedaliodes pheretiades</emphasis>
Grose-Smith &amp; Kirby, 1894
</taxonomicName>
), with the two forming a fully supported clade sister to
<taxonomicName id="589B7982FFF1FFF7FB45D248B4C3821E" box="[1163,1265,1853,1877]" class="Insecta" family="Nymphalidae" genus="Pedaliodes" kingdom="Animalia" order="Lepidoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="genus">
<emphasis id="ADEFDE13FFF1FFF7FB45D248B4C3821E" box="[1163,1265,1853,1877]" italics="true" pageId="5" pageNumber="6">Pedaliodes</emphasis>
</taxonomicName>
+ the monotypic
<taxonomicName id="589B7982FFF1FFF7FCE4D229B497823F" authority="Adams &amp; Bernard, 1979" authorityName="Adams &amp; Bernard" authorityYear="1979" box="[810,1189,1884,1908]" class="Insecta" family="Nymphalidae" genus="Dangond" kingdom="Animalia" order="Lepidoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="genus">
<emphasis id="ADEFDE13FFF1FFF7FCE4D229B3B6823F" box="[810,900,1884,1908]" italics="true" pageId="5" pageNumber="6">Dangond</emphasis>
<bibRefCitation id="FB0A7FF0FFF1FFF7FC5DD229B497823F" author="Adams MJ &amp; Bernard G" box="[915,1189,1884,1908]" pageId="5" pageNumber="6" pagination="95 - 118" refId="ref10911" refString="Adams MJ, Bernard G. Pronophiline buuerflies (Satyridae) of the Serrania de Valledupar, Colombia - Venezuela border. Systematic Entomology 1979; 4: 95 - 118." type="journal article" year="1979">Adams &amp; Bernard, 1979</bibRefCitation>
</taxonomicName>
(
<typeStatus id="4020BCA3FFF1FFF7FB75D229B4DE823F" box="[1211,1260,1884,1908]" pageId="5" pageNumber="6">type</typeStatus>
species:
<taxonomicName id="589B7982FFF1FFF7FA97D229B44982DF" authority="Adams &amp; Bernard, 1979" authorityName="Adams &amp; Bernard" authorityYear="1979" class="Insecta" family="Nymphalidae" genus="Dangond" kingdom="Animalia" order="Lepidoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="species" species="dangondi">
<emphasis id="ADEFDE13FFF1FFF7FA97D229B3B782D8" italics="true" pageId="5" pageNumber="6">Dangond dangondi</emphasis>
<bibRefCitation id="FB0A7FF0FFF1FFF7FC47D20EB44982DF" author="Adams MJ &amp; Bernard G" box="[905,1147,1915,1940]" pageId="5" pageNumber="6" pagination="95 - 118" refId="ref10911" refString="Adams MJ, Bernard G. Pronophiline buuerflies (Satyridae) of the Serrania de Valledupar, Colombia - Venezuela border. Systematic Entomology 1979; 4: 95 - 118." type="journal article" year="1979">Adams &amp; Bernard, 1979</bibRefCitation>
</taxonomicName>
), within the diverse
<taxonomicName id="589B7982FFF1FFF7FA82D20EB58082D8" box="[1356,1458,1915,1939]" class="Insecta" family="Nymphalidae" genus="Pedaliodes" kingdom="Animalia" order="Lepidoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="genus">
<emphasis id="ADEFDE13FFF1FFF7FA82D20EB58082D8" box="[1356,1458,1915,1939]" italics="true" pageId="5" pageNumber="6">Pedaliodes</emphasis>
</taxonomicName>
clade of the subtribe
<taxonomicName id="589B7982FFF1FFF7FBDBD2EEB4AE82F8" box="[1045,1180,1947,1971]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="subTribe" subTribe="Pronophilina">Pronophilina</taxonomicName>
(
<taxonomicName id="589B7982FFF1FFF7FB62D2EEB50D82F8" box="[1196,1343,1947,1971]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="family">Nymphalidae</taxonomicName>
:
<taxonomicName id="589B7982FFF1FFF7FA82D2EEB58282F8" authorityName="Boisduval" authorityYear="1833" box="[1356,1456,1947,1971]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="subFamily" subFamily="Satyrinae">Satyrinae</taxonomicName>
:
<taxonomicName id="589B7982FFF2FFF4FF4FD223B0E78225" authorityName="Boisduval" authorityYear="1833" box="[129,213,1878,1902]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="tribe" tribe="Satyrini">Satyrini</taxonomicName>
) (
<figureCitation id="07A01E84FFF2FFF4FF22D223B1178225" box="[236,293,1878,1902]" captionStart="Figure 1" captionStartId="4.[129,194,1665,1689]" captionTargetBox="[127,1478,145,1637]" captionTargetId="figure-133@4.[127,1478,145,1637]" captionTargetPageId="4" captionText="Figure 1. Maximum likelihood tree of Nivaliodes and other Pronophilina inferred in IQ-TREE based on target enrichment data and COI." pageId="6" pageNumber="7">Fig. 1</figureCitation>
). The topology of
<taxonomicName id="589B7982FFF2FFF4FE28D223B27E8225" authorityName="Pyrcz &amp; Mahecha-J. &amp; Boyer &amp; Lachowska-Cielik &amp; Cerdeña &amp; Farfán &amp; Garlacz &amp; Lorenc-Brudecka &amp; Bálint &amp; Fåhraeus &amp; Zając-Garlacz &amp; Espeland" authorityYear="2024" box="[486,588,1878,1902]" class="Insecta" family="Boreidae" genus="Nivaliodes" kingdom="Animalia" order="Mecoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="genus">
<emphasis id="ADEFDE13FFF2FFF4FE28D223B27E8225" box="[486,588,1878,1902]" italics="true" pageId="6" pageNumber="7">Nivaliodes</emphasis>
</taxonomicName>
shows a fully supported
<taxonomicName id="589B7982FFF2FFF4FF03D203B1A482C5" authorityName="Pyrcz &amp; Mahecha-J. &amp; Boyer &amp; Lachowska-Cielik &amp; Cerdeña &amp; Farfán &amp; Garlacz &amp; Lorenc-Brudecka &amp; Bálint &amp; Fåhraeus &amp; Zając-Garlacz &amp; Espeland" authorityYear="2024" box="[205,406,1910,1934]" class="Insecta" family="Boreidae" genus="Nivaliodes" kingdom="Animalia" order="Mecoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="species" species="viracocha" status="sp. nov.">
<emphasis id="ADEFDE13FFF2FFF4FF03D203B1A482C5" box="[205,406,1910,1934]" italics="true" pageId="6" pageNumber="7">Nivaliodes viracocha</emphasis>
</taxonomicName>
<taxonomicNameLabel id="B6DC6368FFF2FFF4FE53D203B1D982C6" box="[413,491,1910,1933]" pageId="6" pageNumber="7" rank="species">sp. nov.</taxonomicNameLabel>
sister to
<taxonomicName id="589B7982FFF2FFF4FD85D203B2AD82C6" authorityName="Pyrcz &amp; Mahecha-J. &amp; Boyer &amp; Lachowska-Cielik &amp; Cerdeña &amp; Farfán &amp; Garlacz &amp; Lorenc-Brudecka &amp; Bálint &amp; Fåhraeus &amp; Zając-Garlacz &amp; Espeland" authorityYear="2024" box="[587,671,1910,1933]" class="Insecta" family="Boreidae" genus="Nivaliodes" kingdom="Animalia" order="Mecoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="species" species="puriq" status="sp. nov.">
<emphasis id="ADEFDE13FFF2FFF4FD85D203B2AD82C6" box="[587,671,1910,1933]" italics="true" pageId="6" pageNumber="7">N. puriq</emphasis>
</taxonomicName>
<taxonomicNameLabel id="B6DC6368FFF2FFF4FD68D203B2C682C6" box="[678,756,1910,1933]" pageId="6" pageNumber="7" rank="species">sp. nov.</taxonomicNameLabel>
+
<taxonomicName id="589B7982FFF2FFF4FF4ED2E3B13D82E6" authorityName="Pyrcz &amp; Mahecha-J. &amp; Boyer &amp; Lachowska-Cielik &amp; Cerdeña &amp; Farfán &amp; Garlacz &amp; Lorenc-Brudecka &amp; Bálint &amp; Fåhraeus &amp; Zając-Garlacz &amp; Espeland" authorityYear="2024" box="[128,271,1941,1965]" class="Insecta" family="Boreidae" genus="Nivaliodes" kingdom="Animalia" order="Mecoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="species" species="negrobueno" status="sp. nov.">
<emphasis id="ADEFDE13FFF2FFF4FF4ED2E3B13D82E6" box="[128,271,1941,1965]" italics="true" pageId="6" pageNumber="7">N. negrobueno</emphasis>
</taxonomicName>
<taxonomicNameLabel id="B6DC6368FFF2FFF4FED9D2E0B15482E7" box="[279,358,1941,1964]" pageId="6" pageNumber="7" rank="species">sp. nov.</taxonomicNameLabel>
The latter clade is not entirely resolved, with a monophyletic
<taxonomicName id="589B7982FFF2FFF4FE96D2C0B1D18287" authorityName="Pyrcz &amp; Mahecha-J. &amp; Boyer &amp; Lachowska-Cielik &amp; Cerdeña &amp; Farfán &amp; Garlacz &amp; Lorenc-Brudecka &amp; Bálint &amp; Fåhraeus &amp; Zając-Garlacz &amp; Espeland" authorityYear="2024" box="[344,483,1972,1996]" class="Insecta" family="Boreidae" genus="Nivaliodes" kingdom="Animalia" order="Mecoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="species" species="negrobueno" status="sp. nov.">
<emphasis id="ADEFDE13FFF2FFF4FE96D2C0B1D18287" box="[344,483,1972,1996]" italics="true" pageId="6" pageNumber="7">N. negrobueno</emphasis>
</taxonomicName>
<taxonomicNameLabel id="B6DC6368FFF2FFF4FE26D2C0B2018287" box="[488,563,1973,1996]" pageId="6" pageNumber="7" rank="species">sp. nov.</taxonomicNameLabel>
but a paraphyletic
<taxonomicName id="589B7982FFF2FFF4FD3CD2C0B35C8225" authorityName="Pyrcz &amp; Mahecha-J. &amp; Boyer &amp; Lachowska-Cielik &amp; Cerdeña &amp; Farfán &amp; Garlacz &amp; Lorenc-Brudecka &amp; Bálint &amp; Fåhraeus &amp; Zając-Garlacz &amp; Espeland" authorityYear="2024" box="[754,878,1879,1996]" class="Insecta" family="Boreidae" genus="Nivaliodes" kingdom="Animalia" order="Mecoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="species" species="puriq">
<emphasis id="ADEFDE13FFF2FFF4FD3CD2C0B35C8225" box="[754,878,1879,1996]" italics="true" pageId="6" pageNumber="7">N. puriq</emphasis>
</taxonomicName>
owing to the position of the sample DLC127. The latter comes from a different locality from the remaining
<taxonomicName id="589B7982FFF2FFF4FAF8D203B5B482C6" authorityName="Pyrcz &amp; Mahecha-J. &amp; Boyer &amp; Lachowska-Cielik &amp; Cerdeña &amp; Farfán &amp; Garlacz &amp; Lorenc-Brudecka &amp; Bálint &amp; Fåhraeus &amp; Zając-Garlacz &amp; Espeland" authorityYear="2024" box="[1334,1414,1910,1933]" class="Insecta" family="Boreidae" genus="Nivaliodes" kingdom="Animalia" order="Mecoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="species" species="puriq">
<emphasis id="ADEFDE13FFF2FFF4FAF8D203B5B482C6" box="[1334,1414,1910,1933]" italics="true" pageId="6" pageNumber="7">N. puriq</emphasis>
</taxonomicName>
, from which only
<specimenCount id="899DC988FFF2FFF4FC7DD2E0B47E82E6" box="[947,1100,1941,1965]" count="2" pageId="6" pageNumber="7" type="generic">two specimens</specimenCount>
are available now, and only one was sequenced.
</paragraph>
<caption id="CBE45289FFF2FFF4FF4FD39EB1308277" pageId="6" pageNumber="7" startId="6.[129,194,1771,1795]" targetBox="[266,1345,156,1700]" targetPageId="6" targetType="figure">
<paragraph id="9F240201FFF2FFF4FF4FD39EB1308277" blockId="6.[129,1469,1771,1852]" pageId="6" pageNumber="7">
<emphasis id="ADEFDE13FFF2FFF4FF4FD39EB0EA824F" bold="true" box="[129,216,1771,1796]" pageId="6" pageNumber="7">Figure 3.</emphasis>
Adults.
<taxonomicName id="589B7982FFF2FFF4FEEBD39EB2338248" authorityName="Pyrcz &amp; Mahecha-J. &amp; Boyer &amp; Lachowska-Cielik &amp; Cerdeña &amp; Farfán &amp; Garlacz &amp; Lorenc-Brudecka &amp; Bálint &amp; Fåhraeus &amp; Zając-Garlacz &amp; Espeland" authorityYear="2024" box="[293,513,1771,1795]" class="Insecta" family="Boreidae" genus="Nivaliodes" kingdom="Animalia" order="Mecoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="species" species="negrobueno">
<emphasis id="ADEFDE13FFF2FFF4FEEBD39EB2338248" bold="true" box="[293,513,1771,1795]" italics="true" pageId="6" pageNumber="7">Nivaliodes negrobueno</emphasis>
</taxonomicName>
. A, male holotype, dorsal, Abra Negrobueno. B, idem, ventral. C, male paratype, dorsal, road Huarac Huachay. D, idem, ventral. E, female paratype, dorsal, Abra Negrobueno. F, idem, ventral. G, female paratype, dorsal, road HuaracHuachay. H, idem, ventral.
</paragraph>
</caption>
<paragraph id="9F240201FFF3FFF5FF43D5E5B297840E" blockId="7.[113,762,144,325]" pageId="7" pageNumber="8">
The divergence of
<taxonomicName id="589B7982FFF3FFF5FE80D5E5B18685E3" authorityName="Pyrcz &amp; Mahecha-J. &amp; Boyer &amp; Lachowska-Cielik &amp; Cerdeña &amp; Farfán &amp; Garlacz &amp; Lorenc-Brudecka &amp; Bálint &amp; Fåhraeus &amp; Zając-Garlacz &amp; Espeland" authorityYear="2024" box="[334,436,144,168]" class="Insecta" family="Boreidae" genus="Nivaliodes" kingdom="Animalia" order="Mecoptera" pageId="7" pageNumber="8" phylum="Arthropoda" rank="genus">
<emphasis id="ADEFDE13FFF3FFF5FE80D5E5B18685E3" box="[334,436,144,168]" italics="true" pageId="7" pageNumber="8">Nivaliodes</emphasis>
</taxonomicName>
started in the late Pliocene, ~3 Mya, and continued through the Pleistocene. The split between
<taxonomicName id="589B7982FFF3FFF5FFBFD5BAB0E585AC" authorityName="Pyrcz &amp; Mahecha-J. &amp; Boyer &amp; Lachowska-Cielik &amp; Cerdeña &amp; Farfán &amp; Garlacz &amp; Lorenc-Brudecka &amp; Bálint &amp; Fåhraeus &amp; Zając-Garlacz &amp; Espeland" authorityYear="2024" box="[113,215,207,231]" class="Insecta" family="Boreidae" genus="Nivaliodes" kingdom="Animalia" order="Mecoptera" pageId="7" pageNumber="8" phylum="Arthropoda" rank="genus">
<emphasis id="ADEFDE13FFF3FFF5FFBFD5BAB0E585AC" box="[113,215,207,231]" italics="true" pageId="7" pageNumber="8">Nivaliodes</emphasis>
</taxonomicName>
and
<taxonomicName id="589B7982FFF3FFF5FED8D5BAB18085AC" authorityName="Forster" authorityYear="1964" box="[278,434,207,231]" class="Insecta" family="Nymphalidae" genus="Pherepedaliodes" kingdom="Animalia" order="Lepidoptera" pageId="7" pageNumber="8" phylum="Arthropoda" rank="genus">
<emphasis id="ADEFDE13FFF3FFF5FED8D5BAB18085AC" box="[278,434,207,231]" italics="true" pageId="7" pageNumber="8">Pherepedaliodes</emphasis>
</taxonomicName>
occurred at the break of the Miocene and Pliocene some 5 Mya, and was preceded marginally by the separation of the
<emphasis id="ADEFDE13FFF3FFF5FE53D478B20B846E" box="[413,569,269,293]" italics="true" pageId="7" pageNumber="8">Pherepedaliodes</emphasis>
+
<taxonomicName id="589B7982FFF3FFF5FD99D478B28F846E" authorityName="Pyrcz &amp; Mahecha-J. &amp; Boyer &amp; Lachowska-Cielik &amp; Cerdeña &amp; Farfán &amp; Garlacz &amp; Lorenc-Brudecka &amp; Bálint &amp; Fåhraeus &amp; Zając-Garlacz &amp; Espeland" authorityYear="2024" box="[599,701,269,293]" class="Insecta" family="Boreidae" genus="Nivaliodes" kingdom="Animalia" order="Mecoptera" pageId="7" pageNumber="8" phylum="Arthropoda" rank="genus">
<emphasis id="ADEFDE13FFF3FFF5FD99D478B28F846E" box="[599,701,269,293]" italics="true" pageId="7" pageNumber="8">Nivaliodes</emphasis>
</taxonomicName>
clade from
<taxonomicName id="589B7982FFF3FFF5FF67D458B13D840E" box="[169,271,301,325]" class="Insecta" family="Nymphalidae" genus="Pedaliodes" kingdom="Animalia" order="Lepidoptera" pageId="7" pageNumber="8" phylum="Arthropoda" rank="genus">
<emphasis id="ADEFDE13FFF3FFF5FF67D458B13D840E" box="[169,271,301,325]" italics="true" pageId="7" pageNumber="8">Pedaliodes</emphasis>
</taxonomicName>
, which occurred some 5.6 Mya (
<figureCitation id="07A01E84FFF3FFF5FD92D458B2A6840E" box="[604,660,301,325]" captionStart="Figure 2" captionStartId="5.[113,178,1383,1407]" captionTargetBox="[149,1419,146,1351]" captionTargetId="figure-242@5.[146,1426,143,1355]" captionTargetPageId="5" captionText="Figure 2. Divergence analysis of Nivaliodes inferred in BEAST2." pageId="7" pageNumber="8">Fig. 2</figureCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>