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<document id="9445598121EFB10D6552932FBF93213D" ID-CLB-Dataset="38930" ID-DOI="10.11646/zootaxa.4118.1.1" ID-GBIF-Dataset="a4e408c4-2575-4b5c-878b-e1b9010ec619" ID-ISSN="1175-5326" ID-Zenodo-Dep="264597" ID-ZooBank="CFA586DA-10EE-468B-80EE-35351E3845FD" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.taxonomicNames_approvedBy="guilherme" checkinTime="1465242077321" checkinUser="plazi" docAuthor="C, Lees David" docDate="2016" docId="038747324C5AC67A1EB729EDFCB32348" docLanguage="en" docName="zootaxa.4118.pdf.imf" docOrigin="Zootaxa 4118 (1)" docStyle="DocumentStyle:5EBBA59367AD13919D70D935FA04F6A3.19:Zootaxa.2013-.monograph" docStyleId="5EBBA59367AD13919D70D935FA04F6A3" docStyleName="Zootaxa.2013-.monograph" docStyleVersion="19" docTitle="Heteropsis subsimilis C, 2016, sp. nov." docType="treatment" docVersion="12" lastPageNumber="45" masterDocId="FFBE3F4A4C76C6571E202D2DFFD32174" masterDocTitle="Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision" masterLastPageNumber="97" masterPageNumber="1" pageNumber="45" updateTime="1735917000942" updateUser="ExternalLinkService">
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<mods:title id="087CE54124AE4BAF689F151E14E365EF">Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision</mods:title>
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<mods:namePart id="2235BD78140B5DC69A19DB38FBF6F8D0">C, Lees David</mods:namePart>
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<treatment id="038747324C5AC67A1EB729EDFCB32348" ID-DOI="http://doi.org/10.5281/zenodo.6086430" ID-GBIF-Taxon="120817086" ID-Zenodo-Dep="6086430" LSID="urn:lsid:plazi:treatment:038747324C5AC67A1EB729EDFCB32348" httpUri="http://treatment.plazi.org/id/038747324C5AC67A1EB729EDFCB32348" lastPageId="45" lastPageNumber="45" pageId="44" pageNumber="45" scope_family="Nymphalidae" scope_subFamily="Satyrinae" scope_tribe="Satyrini">
<subSubSection id="C334A5AF4C5AC67A1EB729EDFCB32348" lastPageId="45" lastPageNumber="46" pageId="44" pageNumber="45" type="nomenclature">
<paragraph id="8B91F6244C5AC67B1EB729EDFE3A25AF" blockId="44.[151,489,1216,1243]" box="[151,489,1216,1243]" pageId="44" pageNumber="45">
<heading id="D0D941484C5AC67B1EB729EDFE3A25AF" bold="true" box="[151,489,1216,1243]" fontSize="11" level="1" pageId="44" pageNumber="45" reason="1">
<emphasis id="B95A2A364C5AC67B1EB729EDFE3A25AF" bold="true" box="[151,489,1216,1243]" pageId="44" pageNumber="45">
<taxonomicName id="4C2E8DA74C5AC67B1EB729EDFE4425AE" box="[151,407,1216,1242]" class="Insecta" family="Nymphalidae" genus="Heteropsis" kingdom="Animalia" order="Lepidoptera" pageId="44" pageNumber="45" phylum="Arthropoda" rank="species" species="subsimilis">
<emphasis id="B95A2A364C5AC67B1EB729EDFE4425AE" bold="true" box="[151,407,1216,1242]" italics="true" pageId="44" pageNumber="45">Heteropsis subsimilis</emphasis>
</taxonomicName>
group
</emphasis>
</heading>
</paragraph>
<paragraph id="8B91F6244C5AC67A1EB7282AFCB32348" blockId="44.[151,1436,1287,2035]" lastBlockId="45.[151,1437,151,573]" lastPageId="45" lastPageNumber="46" pageId="44" pageNumber="45">
The ‘
<taxonomicName id="4C2E8DA74C5AC67B1EF2282AFE912454" box="[210,322,1287,1312]" class="Insecta" family="Nymphalidae" genus="Heteropsis" kingdom="Animalia" order="Lepidoptera" pageId="44" pageNumber="45" phylum="Arthropoda" rank="species" species="subsimilis">
<emphasis id="B95A2A364C5AC67B1EF2282AFE912454" box="[210,322,1287,1312]" italics="true" pageId="44" pageNumber="45">subsimilis</emphasis>
</taxonomicName>
7’ species group (after
<taxonomicName id="4C2E8DA74C5AC67B1C1B282AFC3E2454" authority="Butler, 1878" authorityName="Butler" authorityYear="1878" box="[571,1005,1287,1312]" class="Insecta" family="Nymphalidae" genus="Pseudonympha" kingdom="Animalia" order="Lepidoptera" pageId="44" pageNumber="45" phylum="Arthropoda" rank="species" species="subsimilis">
<emphasis id="B95A2A364C5AC67B1C1B282AFC8E2454" box="[571,861,1287,1312]" italics="true" pageId="44" pageNumber="45">Pseudonympha subsimilis</emphasis>
Butler, 1878
</taxonomicName>
) of
<bibRefCitation id="EFBF8BD54C5AC67B1A3A2825FB242454" box="[1050,1271,1287,1312]" pageId="44" pageNumber="45" refString="Torres, E., Lees, D. C., Vane-Wright, R. I., Kremen, C., Leonard, J. A. &amp; Wayne, R. K. (2001) Examining monophyly in a large radiation of Madagascan butterflies (Lepidoptera: Satyrinae: Mycalesina) based on mitochondrial DNA data. Molecular Phylogenetics and Evolution, 20, 460 - 473. http: // dx. doi. org / 10.1006 / mpev. 2001.0984" type="journal article">
Torres
<emphasis id="B95A2A364C5AC67B1A482824FB752454" box="[1128,1190,1287,1312]" italics="true" pageId="44" pageNumber="45">et al.,</emphasis>
(2001)
</bibRefCitation>
resemble each other not only in wing pattern, but also in ♂ genitalia. This
<taxonomicName id="4C2E8DA74C5AC67B1D692802FC362433" box="[841,997,1326,1351]" class="Insecta" family="Nymphalidae" genus="Pseudonympha" kingdom="Animalia" order="Lepidoptera" pageId="44" pageNumber="45" phylum="Arthropoda" rank="species" species="subsimilis">
<emphasis id="B95A2A364C5AC67B1D692802FC362433" box="[841,997,1326,1351]" italics="true" pageId="44" pageNumber="45">Ht. subsimilis</emphasis>
</taxonomicName>
group, which occurs within
<collectingCountry id="F339B6B44C5AC67B1B0F2803FA4F2433" box="[1327,1436,1326,1351]" name="Madagascar" pageId="44" pageNumber="45">Malagasy</collectingCountry>
rainforest, has also colonised the
<collectingCountry id="F339B6B44C5AC67B1C01287EFD59241F" box="[545,650,1363,1387]" name="Comoros" pageId="44" pageNumber="45">Comoros</collectingCountry>
, as
<taxonomicName id="4C2E8DA74C5AC67B1C9C287EFBF0241F" authority="Oberthur, 1916" authorityName="Oberthur" authorityYear="1916" box="[700,1059,1362,1387]" class="Insecta" family="Nymphalidae" genus="Culapa" higherTaxonomySource="CoL" kingdom="Animalia" order="Lepidoptera" pageId="44" pageNumber="45" phylum="Arthropoda" rank="species" species="comorana">
<emphasis id="B95A2A364C5AC67B1C9C287EFC86241E" box="[700,853,1363,1386]" italics="true" pageId="44" pageNumber="45">H. comorana</emphasis>
(
<bibRefCitation id="EFBF8BD54C5AC67B1D47287FFBC8241F" author="Oberthur" box="[871,1051,1362,1387]" pageId="44" pageNumber="45" refString="Oberthur, C. (1916) Observations sur une centurie d'Especes de Lepidopteres Rhopaloceres malagaches. Etudes de Lepidopterologie Comparee, 11, 177 - 244." type="journal article" year="1916">Oberthür, 1916</bibRefCitation>
)
</taxonomicName>
, a species closely related to the generally abundant
<taxonomicName id="4C2E8DA74C5AC67B1F56285AFD6524FB" authority="Oberthur, 1916" authorityName="Oberthur" authorityYear="1916" box="[374,694,1398,1423]" class="Insecta" family="Nymphalidae" genus="Culapa" higherTaxonomySource="CoL" kingdom="Animalia" order="Lepidoptera" pageId="44" pageNumber="45" phylum="Arthropoda" rank="species" species="pauper">
<emphasis id="B95A2A364C5AC67B1F56285AFE2324FA" box="[374,496,1399,1422]" italics="true" pageId="44" pageNumber="45">Ht. pauper</emphasis>
(
<bibRefCitation id="EFBF8BD54C5AC67B1C20285BFD7D24FB" author="Oberthur" box="[512,686,1398,1423]" pageId="44" pageNumber="45" refString="Oberthur, C. (1916) Observations sur une centurie d'Especes de Lepidopteres Rhopaloceres malagaches. Etudes de Lepidopterologie Comparee, 11, 177 - 244." type="journal article" year="1916">Oberthür, 1916</bibRefCitation>
)
</taxonomicName>
(
<bibRefCitation id="EFBF8BD54C5AC67B1CE6285AFC9B24FB" author="Lees" box="[710,840,1399,1423]" pageId="44" pageNumber="45" refString="Lees, D. C. (1997) Systematics and biogeography of Madagascan mycalesine butterflies (Lepidoptera: Satyrinae). Unpubl. PhD thesis, University of London, London, 411 pp." type="book" year="1997">Lees, 1997</bibRefCitation>
: 168;
<bibRefCitation id="EFBF8BD54C5AC67B1DAE285AFBB224FB" author="Torres" box="[910,1121,1398,1423]" pageId="44" pageNumber="45" refString="Torres, E., Lees, D. C., Vane-Wright, R. I., Kremen, C., Leonard, J. A. &amp; Wayne, R. K. (2001) Examining monophyly in a large radiation of Madagascan butterflies (Lepidoptera: Satyrinae: Mycalesina) based on mitochondrial DNA data. Molecular Phylogenetics and Evolution, 20, 460 - 473. http: // dx. doi. org / 10.1006 / mpev. 2001.0984" type="journal article" year="2001">
Torres
<emphasis id="B95A2A364C5AC67B1DFD285AFBCE24FB" box="[989,1053,1398,1423]" italics="true" pageId="44" pageNumber="45">et al.,</emphasis>
2001
</bibRefCitation>
). The sister grouping of the
<taxonomicName id="4C2E8DA74C5AC67B1EB728B6FEE224C7" box="[151,305,1434,1459]" class="Insecta" family="Nymphalidae" genus="Pseudonympha" kingdom="Animalia" order="Lepidoptera" pageId="44" pageNumber="45" phylum="Arthropoda" rank="species" species="subsimilis">
<emphasis id="B95A2A364C5AC67B1EB728B6FEE224C7" box="[151,305,1434,1459]" italics="true" pageId="44" pageNumber="45">Ht. subsimilis</emphasis>
</taxonomicName>
group was unresolved in several earlier studies (
<bibRefCitation id="EFBF8BD54C5AC67B1D7528B6FC0624C7" author="Lees" box="[853,981,1435,1459]" pageId="44" pageNumber="45" refString="Lees, D. C. (1997) Systematics and biogeography of Madagascan mycalesine butterflies (Lepidoptera: Satyrinae). Unpubl. PhD thesis, University of London, London, 411 pp." type="book" year="1997">Lees, 1997</bibRefCitation>
,
<bibRefCitation id="EFBF8BD54C5AC67B1DC028B6FB6324C7" author="Torres" box="[992,1200,1434,1459]" pageId="44" pageNumber="45" refString="Torres, E., Lees, D. C., Vane-Wright, R. I., Kremen, C., Leonard, J. A. &amp; Wayne, R. K. (2001) Examining monophyly in a large radiation of Madagascan butterflies (Lepidoptera: Satyrinae: Mycalesina) based on mitochondrial DNA data. Molecular Phylogenetics and Evolution, 20, 460 - 473. http: // dx. doi. org / 10.1006 / mpev. 2001.0984" type="journal article" year="2001">
Torres
<emphasis id="B95A2A364C5AC67B1A0E28B6FBBD24C7" box="[1070,1134,1434,1459]" italics="true" pageId="44" pageNumber="45">et al.,</emphasis>
2001
</bibRefCitation>
,
<bibRefCitation id="EFBF8BD54C5AC67B1A9B28B7FA4B24C7" author="Linares" box="[1211,1432,1434,1459]" pageId="44" pageNumber="45" refString="Linares, M. C., Soto-Calderon, I. D., Lees, D. C. &amp; Anthony, N. M. (2009) High mitochondrial diversity in geographically widespread butterflies of Madagascar: a test of the DNA barcoding approach. Molecular Phylogenetics and Evolution, 50, 485 - 495. http: // dx. doi. org / 10.1016 / j. ympev. 2008.11.008" type="journal article" year="2009">
Linares
<emphasis id="B95A2A364C5AC67B1B3628B6FA9C24C7" box="[1302,1359,1434,1459]" italics="true" pageId="44" pageNumber="45">et al.</emphasis>
, 2009
</bibRefCitation>
,
<bibRefCitation id="EFBF8BD54C5AC67B1EB72893FE3A24A3" author="Kodandaramaiah" box="[151,489,1470,1495]" pageId="44" pageNumber="45" refString="Kodandaramaiah, U., Lees, D. C., Muller, C. J., Torres, E., Karanth, C. P. &amp; Wahlberg, N. (2010) Phylogenetics and biogeography of a spectacular Old World radiation of butterflies: the subtribe Mycalesina (Lepidoptera: Nymphalidae: Satyrini). BMC Evolutionary Biology, 10, 172. http: // dx. doi. org / 10.1186 / 1471 - 2148 - 10 - 172" type="journal article" year="2010">
Kodandaramaiah
<emphasis id="B95A2A364C5AC67B1F402892FE7024A3" box="[352,419,1470,1495]" italics="true" pageId="44" pageNumber="45">et al.,</emphasis>
2010
</bibRefCitation>
), although these studies found good support that the
<taxonomicName id="4C2E8DA74C5AC67B1A772892FB2624A3" box="[1111,1269,1470,1495]" class="Insecta" family="Nymphalidae" genus="Pseudonympha" kingdom="Animalia" order="Lepidoptera" pageId="44" pageNumber="45" phylum="Arthropoda" rank="species" species="subsimilis">
<emphasis id="B95A2A364C5AC67B1A772892FB2624A3" box="[1111,1269,1470,1495]" italics="true" pageId="44" pageNumber="45">Ht. subsimilis</emphasis>
</taxonomicName>
species group forms a clade (97% based on COII parsimony analysis:
<bibRefCitation id="EFBF8BD54C5AC67B1D3828CEFC3F248F" author="Torres" box="[792,1004,1506,1531]" pageId="44" pageNumber="45" refString="Torres, E., Lees, D. C., Vane-Wright, R. I., Kremen, C., Leonard, J. A. &amp; Wayne, R. K. (2001) Examining monophyly in a large radiation of Madagascan butterflies (Lepidoptera: Satyrinae: Mycalesina) based on mitochondrial DNA data. Molecular Phylogenetics and Evolution, 20, 460 - 473. http: // dx. doi. org / 10.1006 / mpev. 2001.0984" type="journal article" year="2001">
Torres
<emphasis id="B95A2A364C5AC67B1D4728CEFC7B248F" box="[871,936,1506,1531]" italics="true" pageId="44" pageNumber="45">et al.,</emphasis>
2001
</bibRefCitation>
; 100% based on maximum likelihood analysis of COI, Ef1a and wingless:
<bibRefCitation id="EFBF8BD54C5AC67B1C602B2BFC42276B" author="Kodandaramaiah" box="[576,913,1542,1567]" pageId="44" pageNumber="45" refString="Kodandaramaiah, U., Lees, D. C., Muller, C. J., Torres, E., Karanth, C. P. &amp; Wahlberg, N. (2010) Phylogenetics and biogeography of a spectacular Old World radiation of butterflies: the subtribe Mycalesina (Lepidoptera: Nymphalidae: Satyrini). BMC Evolutionary Biology, 10, 172. http: // dx. doi. org / 10.1186 / 1471 - 2148 - 10 - 172" type="journal article" year="2010">
Kodandaramaiah
<emphasis id="B95A2A364C5AC67B1D292B2AFC99276B" box="[777,842,1542,1567]" italics="true" pageId="44" pageNumber="45">et al.,</emphasis>
2010
</bibRefCitation>
).
<bibRefCitation id="EFBF8BD54C5AC67B1D862B2BFB4A276B" box="[934,1177,1542,1567]" pageId="44" pageNumber="45" refString="Linares, M. C., Soto-Calderon, I. D., Lees, D. C. &amp; Anthony, N. M. (2009) High mitochondrial diversity in geographically widespread butterflies of Madagascar: a test of the DNA barcoding approach. Molecular Phylogenetics and Evolution, 50, 485 - 495. http: // dx. doi. org / 10.1016 / j. ympev. 2008.11.008" type="journal article">
Linares
<emphasis id="B95A2A364C5AC67B1A232B2AFB96276B" box="[1027,1093,1542,1567]" italics="true" pageId="44" pageNumber="45">et al.,</emphasis>
(2009)
</bibRefCitation>
presented a supported phylogeography of relationships of three species. Recently though, Aduse-Poku
<emphasis id="B95A2A364C5AC67B1A022B06FBB12737" box="[1058,1122,1578,1603]" italics="true" pageId="44" pageNumber="45">et al.,</emphasis>
(2015,
<figureCitation id="1315EAA14C5AC67B1A9A2B07FB2A2737" box="[1210,1273,1578,1603]" captionStart="FIGURE 1" captionStartId="7.[151,250,1711,1733]" captionTargetBox="[166,1422,492,1675]" captionTargetId="figure@7.[150,1437,481,1690]" captionTargetPageId="7" captionText="FIGURE 1. Venation and wing patterning diagram of a fairly typical species of Heteropsis from Madagascar (Heteropis antahala group; ♂), DV. Sdbp = Supra-discocellular brush and patch. Cub = Cubital brush. Ventral banding: Cb = Costal bars. Mf = Median fleck (pale highlight). Mb = Median band. PMb = Premedian band. Ore = Ocellus ring extension. Pml = Premarginal line. Sml = Submarginal line." figureDoi="http://doi.org/10.5281/zenodo.264598" httpUri="https://zenodo.org/record/264598/files/figure.png" pageId="44" pageNumber="45">Fig. 1</figureCitation>
) found strong support for three sampled species namely
<taxonomicName id="4C2E8DA74C5AC67B1C4C2B62FCD62713" box="[620,773,1614,1639]" class="Insecta" family="Nymphalidae" genus="Pseudonympha" kingdom="Animalia" order="Lepidoptera" pageId="44" pageNumber="45" phylum="Arthropoda" rank="species" species="subsimilis">
<emphasis id="B95A2A364C5AC67B1C4C2B62FCD62713" box="[620,773,1614,1639]" italics="true" pageId="44" pageNumber="45">Ht. subsimilis</emphasis>
</taxonomicName>
,
<taxonomicName id="4C2E8DA74C5AC67B1D332B62FC582712" box="[787,907,1615,1638]" class="Insecta" family="Nymphalidae" genus="Culapa" higherTaxonomySource="CoL" kingdom="Animalia" order="Lepidoptera" pageId="44" pageNumber="45" phylum="Arthropoda" rank="species" species="pauper">
<emphasis id="B95A2A364C5AC67B1D332B62FC582712" box="[787,907,1615,1638]" italics="true" pageId="44" pageNumber="45">Ht. pauper</emphasis>
</taxonomicName>
and
<taxonomicName id="4C2E8DA74C5AC67B1DE12B62FB8E2712" box="[961,1117,1615,1638]" class="Insecta" family="Nymphalidae" genus="Culapa" higherTaxonomySource="CoL" kingdom="Animalia" order="Lepidoptera" pageId="44" pageNumber="45" phylum="Arthropoda" rank="species" species="comorana">
<emphasis id="B95A2A364C5AC67B1DE12B62FB8E2712" box="[961,1117,1615,1638]" italics="true" pageId="44" pageNumber="45">Ht. comorana</emphasis>
</taxonomicName>
, which clade they recovered as sister to 13 sampled species of the
<taxonomicName id="4C2E8DA74C5AC67B1C692B5EFD1827FF" box="[585,715,1650,1675]" class="Insecta" family="Nymphalidae" genus="Heteropsis" kingdom="Animalia" order="Lepidoptera" pageId="44" pageNumber="45" phylum="Arthropoda" rank="species" species="strigula">
<emphasis id="B95A2A364C5AC67B1C692B5EFD1827FF" box="[585,715,1650,1675]" italics="true" pageId="44" pageNumber="45">Ht. strigula</emphasis>
</taxonomicName>
group, including
<taxonomicName id="4C2E8DA74C5AC67B1DB92B5EFBCE27FF" box="[921,1053,1650,1675]" class="Insecta" family="Nymphalidae" genus="Heteropsis" kingdom="Animalia" order="Lepidoptera" pageId="44" pageNumber="45" phylum="Arthropoda" rank="species" species="tornado" status="sp. nov.">
<emphasis id="B95A2A364C5AC67B1DB92B5EFBCE27FF" box="[921,1053,1650,1675]" italics="true" pageId="44" pageNumber="45">Ht. tornado</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A269974D4C5AC67B1A062B5FFB5227FF" box="[1062,1153,1650,1675]" pageId="44" pageNumber="45" rank="species">
<emphasis id="B95A2A364C5AC67B1A062B5FFB5227FF" bold="true" box="[1062,1153,1650,1675]" pageId="44" pageNumber="45">sp. nov.</emphasis>
</taxonomicNameLabel>
and the
<taxonomicName id="4C2E8DA74C5AC67B1AC62B5EFAB027FF" box="[1254,1379,1650,1675]" class="Insecta" family="Nymphalidae" genus="Heteropsis" kingdom="Animalia" order="Lepidoptera" pageId="44" pageNumber="45" phylum="Arthropoda" rank="species" species="ankova">
<emphasis id="B95A2A364C5AC67B1AC62B5EFAB027FF" box="[1254,1379,1650,1675]" italics="true" pageId="44" pageNumber="45">Ht. ankova</emphasis>
</taxonomicName>
subgroup (see below). ♂ genitalia have valve tips with a stout inward projecting spine that fall far short of the uncus extension; from LV, the uncus is inflated before its tip. The ♂ genitalia are in fact remarkably miniaturised compared to other
<taxonomicName id="4C2E8DA74C5AC67B1F492BF2FE322782" box="[361,481,1759,1782]" class="Insecta" family="Nymphalidae" genus="Heteropsis" kingdom="Animalia" order="Lepidoptera" pageId="44" pageNumber="45" phylum="Arthropoda" rank="genus">
<emphasis id="B95A2A364C5AC67B1F492BF2FE322782" box="[361,481,1759,1782]" italics="true" pageId="44" pageNumber="45">Heteropsis</emphasis>
</taxonomicName>
, only around
<quantity id="4CD65BC14C5AC67B1C5D2BF2FD012782" box="[637,722,1759,1783]" metricMagnitude="-3" metricUnit="m" metricValue="1.7" pageId="44" pageNumber="45" unit="mm" value="1.7">1.7 mm</quantity>
in length (apart from those of
<taxonomicName id="4C2E8DA74C5AC67B1A082BF2FB782783" box="[1064,1195,1758,1783]" class="Insecta" family="Nymphalidae" genus="Heteropsis" kingdom="Animalia" order="Lepidoptera" pageId="44" pageNumber="45" phylum="Arthropoda" rank="species" species="tornado">
<emphasis id="B95A2A364C5AC67B1A082BF2FB782783" box="[1064,1195,1758,1783]" italics="true" pageId="44" pageNumber="45">Ht. tornado</emphasis>
</taxonomicName>
which is about
<quantity id="4CD65BC14C5AC67B1B7C2BF2FA4F2782" box="[1372,1436,1759,1783]" metricMagnitude="-3" metricUnit="m" metricValue="2.0" pageId="44" pageNumber="45" unit="mm" value="2.0">2 mm</quantity>
long, it can be seen in figures here that those of the
<taxonomicName id="4C2E8DA74C5AC67B1CFD2A2EFC8D266F" box="[733,862,1794,1819]" class="Insecta" family="Nymphalidae" genus="Heteropsis" kingdom="Animalia" order="Lepidoptera" pageId="44" pageNumber="45" phylum="Arthropoda" rank="species" species="strigula">
<emphasis id="B95A2A364C5AC67B1CFD2A2EFC8D266F" box="[733,862,1794,1819]" italics="true" pageId="44" pageNumber="45">Ht. strigula</emphasis>
</taxonomicName>
group are generally between
<quantity id="4CD65BC14C5AC67B1A8C2A2EFAD9266E" box="[1196,1290,1795,1819]" metricMagnitude="-3" metricUnit="m" metricValue="2.5" metricValueMax="3.0" metricValueMin="2.0" pageId="44" pageNumber="45" unit="mm" value="2.5" valueMax="3.0" valueMin="2.0">2–3 mm</quantity>
long and the
<taxonomicName id="4C2E8DA74C5AC67B1EB72A0AFECE264B" box="[151,285,1830,1855]" class="Insecta" family="Nymphalidae" genus="Heteropsis" kingdom="Animalia" order="Lepidoptera" pageId="44" pageNumber="45" phylum="Arthropoda" rank="species" species="drepana">
<emphasis id="B95A2A364C5AC67B1EB72A0AFECE264B" box="[151,285,1830,1855]" italics="true" pageId="44" pageNumber="45">Ht. drepana</emphasis>
</taxonomicName>
and
<taxonomicName id="4C2E8DA74C5AC67B1F722A0AFE33264B" box="[338,480,1830,1855]" class="Insecta" family="Nymphalidae" genus="Mycalesis" kingdom="Animalia" order="Lepidoptera" pageId="44" pageNumber="45" phylum="Arthropoda" rank="species" species="antahala">
<emphasis id="B95A2A364C5AC67B1F722A0AFE33264B" box="[338,480,1830,1855]" italics="true" pageId="44" pageNumber="45">Ht. antahala</emphasis>
</taxonomicName>
groups are more than three mm long and exhibit extension of particular parts). It may be hypothesised that such reduction comes with a relaxation of importance of this character system as a pre-mating isolation mechanism, as apparent in the
<taxonomicName id="4C2E8DA74C5AC67B1C792A42FD2026F3" box="[601,755,1902,1927]" class="Insecta" family="Nymphalidae" genus="Pseudonympha" kingdom="Animalia" order="Lepidoptera" pageId="44" pageNumber="45" phylum="Arthropoda" rank="species" species="subsimilis">
<emphasis id="B95A2A364C5AC67B1C792A42FD2026F3" box="[601,755,1902,1927]" italics="true" pageId="44" pageNumber="45">Ht. subsimilis</emphasis>
</taxonomicName>
group, yet only in the
<taxonomicName id="4C2E8DA74C5AC67B1DD42A42FBA926F3" box="[1012,1146,1902,1927]" class="Insecta" family="Nymphalidae" genus="Heteropsis" kingdom="Animalia" order="Lepidoptera" pageId="44" pageNumber="45" phylum="Arthropoda" rank="species" species="drepana">
<emphasis id="B95A2A364C5AC67B1DD42A42FBA926F3" box="[1012,1146,1902,1927]" italics="true" pageId="44" pageNumber="45">Ht. drepana</emphasis>
</taxonomicName>
group is there a tendency to reduction or even loss (the supra-discocellular patch of
<taxonomicName id="4C2E8DA74C5AC67B1D092ABEFC7126DE" box="[809,930,1939,1962]" class="Insecta" family="Nymphalidae" genus="Heteropsis" kingdom="Animalia" order="Lepidoptera" pageId="44" pageNumber="45" phylum="Arthropoda" rank="species" species="narova">
<emphasis id="B95A2A364C5AC67B1D092ABEFC7126DE" box="[809,930,1939,1962]" italics="true" pageId="44" pageNumber="45">Ht. narova</emphasis>
</taxonomicName>
) of parts of the androconial system (perhaps regained in
<taxonomicName id="4C2E8DA74C5AC67B1F052A9AFE6226BB" box="[293,433,1974,1999]" class="Insecta" family="Nymphalidae" genus="Heteropsis" kingdom="Animalia" order="Lepidoptera" pageId="44" pageNumber="45" phylum="Arthropoda" rank="species" species="drepana">
<emphasis id="B95A2A364C5AC67B1F052A9AFE6226BB" box="[293,433,1974,1999]" italics="true" pageId="44" pageNumber="45">Ht. drepana</emphasis>
</taxonomicName>
only). Atrophication of the genitalic system may be compensated for in a different, possibly androconial character system. Phenotypically, adult ♂♂ and ♀♀ of the
<taxonomicName id="4C2E8DA74C5AC67B1A0F2AF6FB182687" box="[1071,1227,2010,2035]" class="Insecta" family="Nymphalidae" genus="Pseudonympha" kingdom="Animalia" order="Lepidoptera" pageId="44" pageNumber="45" phylum="Arthropoda" rank="species" species="subsimilis">
<emphasis id="B95A2A364C5AC67B1A0F2AF6FB182687" box="[1071,1227,2010,2035]" italics="true" pageId="44" pageNumber="45">Ht. subsimilis</emphasis>
</taxonomicName>
group are similar, nearly monomorphic except in size, but for simple androconia in the ♂ discocellular region and an inflated HW vein system also found in the
<taxonomicName id="4C2E8DA74C5BC67A1FCC2D90FDBD21A1" box="[492,622,188,213]" class="Insecta" family="Nymphalidae" genus="Heteropsis" kingdom="Animalia" order="Lepidoptera" pageId="45" pageNumber="46" phylum="Arthropoda" rank="species" species="strigula">
<emphasis id="B95A2A364C5BC67A1FCC2D90FDBD21A1" box="[492,622,188,213]" italics="true" pageId="45" pageNumber="46">Ht. strigula</emphasis>
</taxonomicName>
group. In the
<taxonomicName id="4C2E8DA74C5BC67A1D302D90FC7F21A1" box="[784,940,188,213]" class="Insecta" family="Nymphalidae" genus="Pseudonympha" kingdom="Animalia" order="Lepidoptera" pageId="45" pageNumber="46" phylum="Arthropoda" rank="species" species="subsimilis">
<emphasis id="B95A2A364C5BC67A1D302D90FC7F21A1" box="[784,940,188,213]" italics="true" pageId="45" pageNumber="46">Ht. subsimilis</emphasis>
</taxonomicName>
group, this system features a set of linearly rather than bulbously inflated veins in the ♂ HW (
<figureCitation id="1315EAA14C5BC67A1CC32DF2FCEA218C" box="[739,825,223,248]" captionStart="FIGURE 12" captionStartId="44.[151,250,1007,1029]" captionTargetBox="[242,1351,208,982]" captionTargetId="figure@44.[212,1370,177,1004]" captionTargetPageId="44" captionText="FIGURE 12. Diagrams of ♂ HW inflated vein system in the Ht. subsimilis group (modified after Lees, 1997: 96); wings in DV and R vein component on the anterior border of the cell in VV (flipped). A: Ht. pauper Oberthür, 1916 (Ankazomivady; “ H. 23 ” in Torres et al., 2001). B: Ht. pauper (Anjanaharibe Sud). C: Ht. kremenae Lees, sp. nov. (Masoala). D: Ht. subsimilis (Butler, 1878) (Ambohitantely), showing M 1 + M 2 inflation and asymmetric swelling above cell. E: Ht. avaratra Lees &amp; Kremen, sp. nov. (Montagne d’Ambre). No wing preparation was available for Ht. sogai." figureDoi="http://doi.org/10.5281/zenodo.264609" httpUri="https://zenodo.org/record/264609/files/figure.png" pageId="45" pageNumber="46">Fig. 12</figureCitation>
;
<bibRefCitation id="EFBF8BD54C5BC67A1D682DCDFC1A218C" author="Lees" box="[840,969,224,248]" pageId="45" pageNumber="46" refString="Lees, D. C. (1997) Systematics and biogeography of Madagascan mycalesine butterflies (Lepidoptera: Satyrinae). Unpubl. PhD thesis, University of London, London, 411 pp." type="book" year="1997">Lees, 1997</bibRefCitation>
: 96). As predominant in the
<taxonomicName id="4C2E8DA74C5BC67A1B382DCCFA4F218C" box="[1304,1436,223,248]" class="Insecta" family="Nymphalidae" genus="Heteropsis" kingdom="Animalia" order="Lepidoptera" pageId="45" pageNumber="46" phylum="Arthropoda" rank="species" species="strigula">
<emphasis id="B95A2A364C5BC67A1B382DCCFA4F218C" box="[1304,1436,223,248]" italics="true" pageId="45" pageNumber="46">Ht. strigula</emphasis>
</taxonomicName>
group, all species have the reduced ocellus configuration of the HWD, with only that in space CuA1 expressed as well as often that in space Rs; the morphological ‘total evidence’ phylogenetic hypotheses of
<bibRefCitation id="EFBF8BD54C5BC67A1A8C2C05FAB82034" author="Lees" box="[1196,1387,295,320]" pageId="45" pageNumber="46" refString="Lees, D. C. (1997) Systematics and biogeography of Madagascan mycalesine butterflies (Lepidoptera: Satyrinae). Unpubl. PhD thesis, University of London, London, 411 pp." type="book" year="1997">Lees (1997: 156)</bibRefCitation>
that the
<taxonomicName id="4C2E8DA74C5BC67A1EE12C60FE8F2011" box="[193,348,332,357]" class="Insecta" family="Nymphalidae" genus="Pseudonympha" kingdom="Animalia" order="Lepidoptera" pageId="45" pageNumber="46" phylum="Arthropoda" rank="species" species="subsimilis">
<emphasis id="B95A2A364C5BC67A1EE12C60FE8F2011" box="[193,348,332,357]" italics="true" pageId="45" pageNumber="46">Ht. subsimilis</emphasis>
</taxonomicName>
group is sister to only part of the
<taxonomicName id="4C2E8DA74C5BC67A1CC02C60FCB12011" box="[736,866,332,357]" class="Insecta" family="Nymphalidae" genus="Heteropsis" kingdom="Animalia" order="Lepidoptera" pageId="45" pageNumber="46" phylum="Arthropoda" rank="species" species="strigula">
<emphasis id="B95A2A364C5BC67A1CC02C60FCB12011" box="[736,866,332,357]" italics="true" pageId="45" pageNumber="46">Ht. strigula</emphasis>
</taxonomicName>
group (see also
<bibRefCitation id="EFBF8BD54C5BC67A1A3E2C60FB3D2010" author="Torres" box="[1054,1262,332,357]" pageId="45" pageNumber="46" refString="Torres, E., Lees, D. C., Vane-Wright, R. I., Kremen, C., Leonard, J. A. &amp; Wayne, R. K. (2001) Examining monophyly in a large radiation of Madagascan butterflies (Lepidoptera: Satyrinae: Mycalesina) based on mitochondrial DNA data. Molecular Phylogenetics and Evolution, 20, 460 - 473. http: // dx. doi. org / 10.1006 / mpev. 2001.0984" type="journal article" year="2001">
Torres
<emphasis id="B95A2A364C5BC67A1A4D2C60FB7E2011" box="[1133,1197,332,357]" italics="true" pageId="45" pageNumber="46">et al.,</emphasis>
2001
</bibRefCitation>
) is presumably biased by the inclusion of this character system, whereas the unsupported Bayesian topology in
<bibRefCitation id="EFBF8BD54C5BC67A1AE22C42FED420D8" author="Kodandaramaiah" pageId="45" pageNumber="46" refString="Kodandaramaiah, U., Lees, D. C., Muller, C. J., Torres, E., Karanth, C. P. &amp; Wahlberg, N. (2010) Phylogenetics and biogeography of a spectacular Old World radiation of butterflies: the subtribe Mycalesina (Lepidoptera: Nymphalidae: Satyrini). BMC Evolutionary Biology, 10, 172. http: // dx. doi. org / 10.1186 / 1471 - 2148 - 10 - 172" type="journal article" year="2010">
Kodandaramaiah
<emphasis id="B95A2A364C5BC67A1BA82C5CFF6820D9" italics="true" pageId="45" pageNumber="46">et al.,</emphasis>
(2010
</bibRefCitation>
,
<figureCitation id="1315EAA14C5BC67A1F342CB9FE8420D8" box="[276,343,404,429]" captionStart="FIGURE 3" captionStartId="15.[151,250,1650,1672]" captionTargetBox="[303,1292,195,1608]" captionTargetId="figure@15.[275,1307,177,1626]" captionTargetPageId="15" captionText="FIGURE 3. Adults in DV (left), VV (right) unless otherwise stated. A: depictions (‘ iconotypes’) of Gallienia alaokola Oberthür (1916) ♂, left (VV) and ♀, right (DV), representing Oberthür’s preference for the type specimens of this nominal species, both referred here to Ht. mabillei (Butler, 1879). B: designated LT ♂ of G. alaokola Oberthür, 1916 (‘ Fianarantsoa’; BMNH (E) 1717102). C: HT ♂ of Strabena mabillei Butler, 1879 (BMNH (E) 672624). Scale bar: 30 mm. D: ♂ abdomen of Oberthür collection specimen of Ht. mabillei (left, VV). E: PLT ♂ abdomen of G. alaokola (Fianarantsoa; BMNH (E) 672621) (right, VV)." figureDoi="http://doi.org/10.5281/zenodo.264600" httpUri="https://zenodo.org/record/264600/files/figure.png" pageId="45" pageNumber="46">Fig. 3</figureCitation>
) would imply that this configuration of HWD reduction is convergent between these two sister species groups (only with a gain of expression in HWD space-M
<quantity id="4CD65BC14C5BC67A1D542C95FC4E20A4" box="[884,925,439,464]" metricMagnitude="-2" metricUnit="m" metricValue="7.62" pageId="45" pageNumber="46" unit="in" value="3.0">3 in</quantity>
<taxonomicName id="4C2E8DA74C5BC67A1D842C94FB9220A4" box="[932,1089,441,464]" class="Insecta" family="Nymphalidae" genus="Heteropsis" kingdom="Animalia" order="Lepidoptera" pageId="45" pageNumber="46" phylum="Arthropoda" rank="species" species="roussettae" status="sp. nov.">
<emphasis id="B95A2A364C5BC67A1D842C94FB9220A4" box="[932,1089,441,464]" italics="true" pageId="45" pageNumber="46">Ht. roussettae</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A269974D4C5BC67A1A682C9AFB7020A4" box="[1096,1187,439,464]" pageId="45" pageNumber="46" rank="species">
<emphasis id="B95A2A364C5BC67A1A682C9AFB7020A4" bold="true" box="[1096,1187,439,464]" pageId="45" pageNumber="46">sp. nov.</emphasis>
</taxonomicNameLabel>
). In any case, the two groups are sisters (
<bibRefCitation id="EFBF8BD54C5BC67A1F4F2CF1FD582081" author="Aduse-Poku" box="[367,651,476,501]" pageId="45" pageNumber="46" refString="Aduse-Poku, K., Lees, D. C., Brattstrom, O., Kodandaramaiah, U., Collins, S. &amp; Brakefield, P. M., (2016) Molecular phylogeny and generic-level taxonomy of the widespread palaeotropical ' Heteropsis clade' (Nymphalidae: Satyrinae: Mycalesina). Systematic Entomology, in press. http: // dx. doi. org / syen. 12183" type="book" year="2016">
Aduse-Poku
<emphasis id="B95A2A364C5BC67A1C252CF0FD932081" box="[517,576,476,501]" italics="true" pageId="45" pageNumber="46">et al.</emphasis>
, 2016
</bibRefCitation>
, in press). Both sexes are attracted to ripe fruit as adults, as well as sometimes other sugar sources, and these low flying butterflies seem to specialise on grasses in the forest interior. There seem to be ridge and riparian specialists (
<bibRefCitation id="EFBF8BD54C5BC67A1C8E2F09FC802348" author="Kremen" box="[686,851,548,572]" pageId="45" pageNumber="46" refString="Kremen, C. (1994) Biological inventory using target taxa: a case study of the butterflies of Madagascar. Ecological Applications, 4, 407 - 422. http: // dx. doi. org / 10.2307 / 1941946" type="journal article" year="1994">Kremen, 1994</bibRefCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>
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