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<mods:title id="53A658F82924927DCE7E9F996F63F84B">First representatives of the stonefly genus Perlomyia and the beaded lacewing genus Isoscelipteron from the Upper Miocene of France (Plecoptera: Leuctridae and Neuroptera: Berothidae), with biogeographical considerations</mods:title>
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<mods:namePart id="2208E1CC80B5ADC2EF9852ACF10AD2B2">Boderau, Mathieu</mods:namePart>
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<mods:namePart id="D6F69AB13BA7613FE2E2BB9125058B26">Ngo-Muller, Valerie</mods:namePart>
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<heading id="1E4F8189B000FFA3F4E16A166A3D8207" bold="true" box="[151,541,1050,1077]" fontSize="11" level="1" pageId="5" pageNumber="136" reason="1">
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<taxonomicName id="82B84D66B000FFA3F4E16A1669988207" authorityName="Boderau &amp; Ngo-Muller &amp; Nel" authorityYear="2024" box="[151,440,1050,1077]" class="Insecta" family="Berothidae" genus="Isoscelipteron" kingdom="Animalia" order="Neuroptera" pageId="5" pageNumber="136" phylum="Arthropoda" rank="species" species="bertrandi" status="sp. nov.">
<emphasis id="77CCEAF7B000FFA3F4E16A1669988207" bold="true" box="[151,440,1050,1077]" italics="true" pageId="5" pageNumber="136">Isoscelipteron bertrandi</emphasis>
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<taxonomicNameLabel id="6CFF578CB000FFA3F5B66A176A3D8207" box="[448,541,1051,1077]" pageId="5" pageNumber="136" rank="species">sp. nov.</taxonomicNameLabel>
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<figureCitation id="DD832A60B000FFA3F4E16A6F68F9824F" box="[151,217,1122,1149]" captionStart="FIGURE 5" captionStartId="6.[152,255,758,782]" captionTargetBox="[151,1435,181,734]" captionTargetId="figure-30@6.[151,1435,181,734]" captionTargetPageId="6" captionText="FIGURE 5. Isoscelipteron bertrandi sp. nov., holotype IF-STR-0105. Photograph of forewing. Scale bar = 2 mm." figureDoi="http://doi.org/10.5281/zenodo.12742786" httpUri="https://zenodo.org/record/12742786/files/figure.png" pageId="5" pageNumber="136">Fig. 5</figureCitation>
</paragraph>
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<subSubSection id="0DA2656EB000FFA3F4E16AA6691182DB" pageId="5" pageNumber="136" type="materials_examined">
<paragraph id="450736E5B000FFA3F4E16AA6691182DB" blockId="5.[151,1437,1194,2013]" pageId="5" pageNumber="136">
<emphasis id="77CCEAF7B000FFA3F4E16AA6692682F6" bold="true" box="[151,262,1194,1220]" pageId="5" pageNumber="136">Material.</emphasis>
<materialsCitation id="F5D03CB8B000FFA3F57B6AA7691182DB" country="France" location="Museum National d'Histoire Naturelle" municipality="Paris" pageId="5" pageNumber="136" specimenCount="1" typeStatus="holotype">
<typeStatus id="9A038847B000FFA3F57B6AA7695582F7" box="[269,373,1195,1221]" pageId="5" pageNumber="136" type="holotype">Holotype</typeStatus>
IF-STR-0105 (a forewing), provisionally housed in the
<location id="4067603EB000FFA3F7906AA76DB882F7" LSID="urn:lsid:plazi:treatment:CD1187F3B000FFA0F4E16A166AB985EB:4067603EB000FFA3F7906AA76DB882F7" box="[998,1432,1195,1221]" country="France" municipality="Paris" name="Museum National d'Histoire Naturelle" pageId="5" pageNumber="136">Muséum National dHistoire Naturelle</location>
,
<collectingMunicipality id="A563AC9FB000FFA3F4E16AC368F282DB" box="[151,210,1231,1257]" pageId="5" pageNumber="136">Paris</collectingMunicipality>
,
<collectingCountry id="3DAF7675B000FFA3F4AB6AC3690D82DB" box="[221,301,1231,1257]" name="France" pageId="5" pageNumber="136">France</collectingCountry>
.
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</paragraph>
</subSubSection>
<subSubSection id="0DA2656EB000FFA3F4B16AFF6A0F834B" pageId="5" pageNumber="136" type="etymology">
<paragraph id="450736E5B000FFA3F4B16AFF69828303" blockId="5.[151,1437,1194,2013]" pageId="5" pageNumber="136">
<emphasis id="77CCEAF7B000FFA3F4B16AFF6968833F" bold="true" box="[199,328,1267,1293]" pageId="5" pageNumber="136">Etymology</emphasis>
. Named after Mr Guilhem Bertrand, for the great help he provided in our field research in the Quarry of Sainte Reine.
</paragraph>
<paragraph id="450736E5B000FFA3F4B16B366A0F834B" blockId="5.[151,1437,1194,2013]" pageId="5" pageNumber="136">
<emphasis id="77CCEAF7B000FFA3F4B16B3669E68366" bold="true" box="[199,454,1338,1365]" pageId="5" pageNumber="136">Locality and horizon</emphasis>
. Latest Miocene, diatom paleomaar, Sainte-Reine (quarry of Foufouilloux, Virargues village), near Murat, Cantal,
<collectingCountry id="3DAF7675B000FFA3F5AD6B536A0B834B" box="[475,555,1375,1401]" name="France" pageId="5" pageNumber="136">France</collectingCountry>
.
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<paragraph id="450736E5B000FFA3F4B16B8E6AFC83D7" blockId="5.[151,1437,1194,2013]" pageId="5" pageNumber="136">
<emphasis id="77CCEAF7B000FFA3F4B16B8E696283AE" bold="true" box="[199,322,1410,1436]" pageId="5" pageNumber="136">Diagnosis.</emphasis>
Forewing characters only. Concavity of postero-distal margin present but not strongly pronounced; 10 posterior branches of RP; line of gradate crossveins making an angle of ca. 20° with the main stem of RP; CuA and CuP with numerous very long parallel branches
</paragraph>
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<emphasis id="77CCEAF7B000FFA3F4B16BE26979803A" bold="true" box="[199,345,1518,1544]" pageId="5" pageNumber="136">Description.</emphasis>
A nearly complete forewing, with area between C and R/RA missing; membrane brow, main veins white, and gradate crossvein black; wing
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long, ca.
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wide; apex of wing pointed; postero-distal margin weakly but distinctly falcate; RP with 10 posterior branches; inner gradate series of crossveins complete, in a weakly curved line making an angle of ca. 20° with the main stem of RP; no outer gradate series of crossveins; MP with two main forks and several long apical branches; CuA and CuP with numerous long parallel branches.
</paragraph>
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<paragraph id="450736E5B000FFA3F4B168AF6BFA81EF" blockId="5.[151,1437,1194,2013]" pageId="5" pageNumber="136">
<emphasis id="77CCEAF7B000FFA3F4B168AF6909808F" bold="true" box="[199,297,1699,1725]" pageId="5" pageNumber="136">Remark</emphasis>
. The acute wing apex, the falcate postero-distal margin of this forewing plus the wing densely covered with setae support an attribution to the
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: Berothinae. Within this subfamily, only some species of the genus
<taxonomicName id="82B84D66B000FFA3F49668E7695E8137" authorityName="Costa" authorityYear="1863" box="[224,382,1771,1797]" class="Insecta" family="Berothidae" genus="Isoscelipteron" kingdom="Animalia" order="Neuroptera" pageId="5" pageNumber="136" phylum="Arthropoda" rank="genus">
<emphasis id="77CCEAF7B000FFA3F49668E7695E8137" box="[224,382,1771,1797]" italics="true" pageId="5" pageNumber="136">Isoscelipteron</emphasis>
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have 10 branches of RP or more, as in the new fossil (Navás 1929;
<bibRefCitation id="21294B14B000FFA3F00E68E76DB78137" author="MacLeod, E. G. &amp; Adams, A." box="[1144,1431,1771,1797]" pageId="5" pageNumber="136" pagination="237 - 265" refId="ref5646" refString="MacLeod, E. G. &amp; Adams, A. (1967) A review of the taxonomy and morphology of the Berothidae with the description of a new subfamily from Chile. Psyche, 74, 237 - 265. https: // doi. org / 10.1155 / 1967 / 898204" type="journal article" year="1967">MacLeod &amp; Adams 1967</bibRefCitation>
;
<bibRefCitation id="21294B14B000FFA3F4E169036983811B" author="Aspock, U. &amp; Aspock, H." box="[151,419,1807,1833]" pageId="5" pageNumber="136" pagination="1 - 14" refId="ref3661" refString="Aspock, U. &amp; Aspock, H. (1981) Weitere Untersuchungen an Berothiden: Berotha Walker, Isoscelipteron Costa und Asadeteva n. g. (Neuropteroidea: Planipennia). Zeitschrift der Arbeitsgemeinschaft Osterreich Entomologen, 33, 1 - 14." type="journal article" year="1981">Aspöck &amp; Aspöck 1981</bibRefCitation>
,
<bibRefCitation id="21294B14B000FFA3F5D9690369D7811B" author="Aspock, U. &amp; Aspock, H." box="[431,503,1807,1833]" pageId="5" pageNumber="136" pagination="65 - 83" refId="ref3707" refString="Aspock, U. &amp; Aspock, H. (1984 a) Zur Kenntnis des Genus Sphaeroberotha Navas, 1930 (Neuropteroidea: Planipennia: Berothidae). Zeitschrift der Arbeitsgemeinschaft Osterreich Entomologen, 35, 65 - 83." type="journal article" year="1984">1984a</bibRefCitation>
,b, 1985a,b, 1986, 1987, 1988a,b,c, 1989, 1991;
<bibRefCitation id="21294B14B000FFA3F07A69036C91811B" author="Smithers, C. N." box="[1036,1201,1807,1833]" pageId="5" pageNumber="136" pagination="4 - 5" refId="ref6719" refString="Smithers, C. N. (1984) The Neuroptera of Barrow and nearby islands off the west coast of Western Australia. Australian Entomological Magazine, 11 (4 - 5), 61 - 68." type="journal article" year="1984">Smithers 1984</bibRefCitation>
;
<bibRefCitation id="21294B14B000FFA3F0CD69036D6B811B" author="Adams, P. A." box="[1211,1355,1807,1833]" pageId="5" pageNumber="136" pagination="187 - 193" refId="ref3576" refString="Adams, P. A. (1989) A new genus of Berothidae from tropical America, with two new species. Psyche, 96 187 - 193. https: // doi. org / 10.1155 / 1989 / 39647" type="journal article" year="1989">Adams 1989</bibRefCitation>
;
<bibRefCitation id="21294B14B000FFA3F12069036945817F" author="Penny, N. D. &amp; Winterton, S. L." pageId="5" pageNumber="136" pagination="1 - 6" refId="ref6276" refString="Penny, N. D. &amp; Winterton, S. L. (2007) Rediscovery of the unusual genus Ormiscocerus (Neuroptera: Berothidae: Cyrenoberothinae). Proceedings of the California Academy of Sciences, (4), 58, 1 - 6." type="journal article" year="2007">Penny &amp; Winterton 2007</bibRefCitation>
;
<bibRefCitation id="21294B14B000FFA3F506693F6A70817F" author="Machado, R. J. P. &amp; Martins, C. C. &amp; Aspock, H. &amp; De Miranda Tavares, L. G. &amp; Aspock, U." box="[368,592,1843,1869]" pageId="5" pageNumber="136" pagination="1422 - 1444" refId="ref5561" refString="Machado, R. J. P., Martins, C. C., Aspock, H., De Miranda Tavares, L. G. &amp; Aspock, U. (2022) The first cave associated genus of Berothidae (Insecta: Neuroptera), and a new interpretation of the subfamily Cyrenoberothinae. Zoological Journal of the Linnean Society, 195, 1422 - 1444. https: // doi. org / 10.1093 / zoolinnean / zlab 104" type="journal article" year="2022">Machado et al 2022</bibRefCitation>
). These are
<taxonomicName id="82B84D66B000FFA3F6A5693F6CD3817F" authority="Aspock &amp; Aspock, 1991" authorityName="Aspock &amp; Aspock" authorityYear="1991" box="[723,1267,1843,1869]" class="Insecta" family="Berothidae" genus="Isoscelipteron" kingdom="Animalia" order="Neuroptera" pageId="5" pageNumber="136" phylum="Arthropoda" rank="species" species="borneole">
<emphasis id="77CCEAF7B000FFA3F6A5693F6BFB817F" box="[723,987,1843,1869]" italics="true" pageId="5" pageNumber="136">Isoscelipteron borneole</emphasis>
<bibRefCitation id="21294B14B000FFA3F796693F6CD3817F" author="Aspock, U. &amp; Aspock, H." box="[992,1267,1843,1869]" pageId="5" pageNumber="136" pagination="65 - 76" refId="ref4196" refString="Aspock, U. &amp; Aspock, H. (1991) Zur Kenntnis des Genus Isoscelipteron Costa, 1863 (Berothidae: Berothinae). Zeitschrift der Arbeitsgemeinschaft Osterreich Entomologen, 43, 65 - 76." type="journal article" year="1991">Aspöck &amp; Aspöck, 1991</bibRefCitation>
</taxonomicName>
,
<taxonomicName id="82B84D66B000FFA3F089693F6A758143" authority="Aspock &amp; Aspock, 1991" authorityName="Aspock &amp; Aspock" authorityYear="1991" class="Insecta" family="Berothidae" genus="Isoscelipteron" kingdom="Animalia" order="Neuroptera" pageId="5" pageNumber="136" phylum="Arthropoda" rank="species" species="philippinicola">
<emphasis id="77CCEAF7B000FFA3F089693F69148143" italics="true" pageId="5" pageNumber="136">Isoscelipteron philippinicola</emphasis>
<bibRefCitation id="21294B14B000FFA3F54D695B6A758143" author="Aspock, U. &amp; Aspock, H." box="[315,597,1879,1905]" pageId="5" pageNumber="136" pagination="65 - 76" refId="ref4196" refString="Aspock, U. &amp; Aspock, H. (1991) Zur Kenntnis des Genus Isoscelipteron Costa, 1863 (Berothidae: Berothinae). Zeitschrift der Arbeitsgemeinschaft Osterreich Entomologen, 43, 65 - 76." type="journal article" year="1991">Aspöck &amp; Aspöck, 1991</bibRefCitation>
</taxonomicName>
, and
<taxonomicName id="82B84D66B000FFA3F6E5695B6C358143" authority="Costa, 1863" authorityName="Costa" authorityYear="1863" box="[659,1045,1879,1905]" class="Insecta" family="Berothidae" genus="Isoscelipteron" kingdom="Animalia" order="Neuroptera" pageId="5" pageNumber="136" phylum="Arthropoda" rank="species" species="fulvum">
<emphasis id="77CCEAF7B000FFA3F6E5695B6BA58143" box="[659,901,1879,1905]" italics="true" pageId="5" pageNumber="136">Isoscelipteron fulvum</emphasis>
<bibRefCitation id="21294B14B000FFA3F7F8695B6C358143" author="Costa, A." box="[910,1045,1879,1905]" pageId="5" pageNumber="136" pagination="1 - 80" refId="ref4919" refString="Costa, A. (1863) Nuovi studii sulla entomologia della Calabria ulteriore. Atti della Accademia delle Scienze Fisiche e Matematiche di Napoli, 1, 1 - 80. https: // doi. org / 10.5962 / bhl. title. 34236" type="journal article" year="1863">Costa, 1863</bibRefCitation>
</taxonomicName>
(
<bibRefCitation id="21294B14B000FFA3F053695B6CE08143" author="Aspock, U. &amp; Aspock, H." box="[1061,1216,1879,1905]" pageId="5" pageNumber="136" pagination="89 - 102" refId="ref3933" refString="Aspock, U. &amp; Aspock, H. (1987) Die Berothiden Australiens. 4: Weitere Untersuchungen uber das Genus Spermophorella Tillyard (Neuropteroidea: Planipennia: Berothidae). Zeitschrift der Arbeitsgemeinschaft Osterreich Entomologen, 38, 89 - 102." type="journal article" year="1987">Aspöck 1987</bibRefCitation>
;
<bibRefCitation id="21294B14B000FFA3F0BA695B68F281A7" author="Aspock, U. &amp; Aspock, H." pageId="5" pageNumber="136" pagination="65 - 76" refId="ref4196" refString="Aspock, U. &amp; Aspock, H. (1991) Zur Kenntnis des Genus Isoscelipteron Costa, 1863 (Berothidae: Berothinae). Zeitschrift der Arbeitsgemeinschaft Osterreich Entomologen, 43, 65 - 76." type="journal article" year="1991">Aspöck &amp; Aspöck 1991</bibRefCitation>
), while the other species have at most eight branches of RP. The new fossil and
<taxonomicName id="82B84D66B000FFA3F01269776D7681A7" authorityName="Costa" authorityYear="1863" box="[1124,1366,1915,1941]" class="Insecta" family="Berothidae" genus="Isoscelipteron" kingdom="Animalia" order="Neuroptera" pageId="5" pageNumber="136" phylum="Arthropoda" rank="species" species="fulvum">
<emphasis id="77CCEAF7B000FFA3F01269776D7681A7" box="[1124,1366,1915,1941]" italics="true" pageId="5" pageNumber="136">Isoscelipteron fulvum</emphasis>
</taxonomicName>
differ from
<taxonomicName id="82B84D66B000FFA3F4AF699369C6818B" authorityName="Aspock &amp; Aspock" authorityYear="1991" box="[217,486,1951,1977]" class="Insecta" family="Berothidae" genus="Isoscelipteron" kingdom="Animalia" order="Neuroptera" pageId="5" pageNumber="136" phylum="Arthropoda" rank="species" species="borneole">
<emphasis id="77CCEAF7B000FFA3F4AF699369C6818B" box="[217,486,1951,1977]" italics="true" pageId="5" pageNumber="136">Isoscelipteron borneole</emphasis>
</taxonomicName>
and
<taxonomicName id="82B84D66B000FFA3F65269936B4A818B" authorityName="Aspock &amp; Aspock" authorityYear="1991" box="[548,874,1951,1977]" class="Insecta" family="Berothidae" genus="Isoscelipteron" kingdom="Animalia" order="Neuroptera" pageId="5" pageNumber="136" phylum="Arthropoda" rank="species" species="philippinicola">
<emphasis id="77CCEAF7B000FFA3F65269936B4A818B" box="[548,874,1951,1977]" italics="true" pageId="5" pageNumber="136">Isoscelipteron philippinicola</emphasis>
</taxonomicName>
in the clearly less pronounced concavity of the postero-distal margin of their forewings (
<bibRefCitation id="21294B14B000FFA3F61569CF6B5681EF" author="Aspock, U. &amp; Aspock, H." box="[611,886,1987,2013]" pageId="5" pageNumber="136" pagination="65 - 76" refId="ref4196" refString="Aspock, U. &amp; Aspock, H. (1991) Zur Kenntnis des Genus Isoscelipteron Costa, 1863 (Berothidae: Berothinae). Zeitschrift der Arbeitsgemeinschaft Osterreich Entomologen, 43, 65 - 76." type="journal article" year="1991">Aspöck &amp; Aspöck 1991</bibRefCitation>
: fig. 12).
</paragraph>
<caption id="11C7666DB003FFA0F4EE6CFA6D04853F" ID-DOI="http://doi.org/10.5281/zenodo.12742786" ID-Zenodo-Dep="12742786" box="[152,1316,758,782]" httpUri="https://zenodo.org/record/12742786/files/figure.png" pageId="6" pageNumber="137" startId="6.[152,255,758,782]" targetBox="[151,1435,181,734]" targetPageId="6" targetType="figure">
<paragraph id="450736E5B003FFA0F4EE6CFA6D04853F" blockId="6.[152,1316,758,782]" box="[152,1316,758,782]" pageId="6" pageNumber="137">
<emphasis id="77CCEAF7B003FFA0F4EE6CFA6938853C" bold="true" box="[152,280,758,782]" pageId="6" pageNumber="137">FIGURE 5.</emphasis>
<taxonomicName id="82B84D66B003FFA0F5696CFA6A34853C" authorityName="Boderau &amp; Ngo-Muller &amp; Nel" authorityYear="2024" box="[287,532,758,782]" class="Insecta" family="Berothidae" genus="Isoscelipteron" kingdom="Animalia" order="Neuroptera" pageId="6" pageNumber="137" phylum="Arthropoda" rank="species" species="bertrandi" status="sp. nov.">
<emphasis id="77CCEAF7B003FFA0F5696CFA6A34853C" box="[287,532,758,782]" italics="true" pageId="6" pageNumber="137">Isoscelipteron bertrandi</emphasis>
</taxonomicName>
<emphasis id="77CCEAF7B003FFA0F66C6CFA6A4A853C" bold="true" box="[538,618,758,782]" pageId="6" pageNumber="137">
<taxonomicNameLabel id="6CFF578CB003FFA0F66C6CFA6A4A853C" box="[538,618,758,782]" pageId="6" pageNumber="137" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
, holotype IF-STR-0105. Photograph of forewing. Scale bar = 2 mm.
</paragraph>
</caption>
<paragraph id="450736E5B003FFA0F4B16D5F6AB985EB" blockId="6.[151,1436,851,985]" pageId="6" pageNumber="137">
The new fossil differs from
<taxonomicName id="82B84D66B003FFA0F67E6D586A4D855F" authorityName="Costa" authorityYear="1863" box="[520,621,851,877]" class="Insecta" family="Berothidae" genus="Isoscelipteron" kingdom="Animalia" order="Neuroptera" pageId="6" pageNumber="137" phylum="Arthropoda" rank="species" species="fulvum">
<emphasis id="77CCEAF7B003FFA0F67E6D586A4D855F" box="[520,621,851,877]" italics="true" pageId="6" pageNumber="137">I. fulvum</emphasis>
</taxonomicName>
in the line of gradating crossveins making an angle of ca. 20° with the main stem of RP vs. ca. 40° in the latter. Additionally, CuA and CuP have numerous very long parallel branches, similar to the situation in
<taxonomicName id="82B84D66B003FFA0F5C76D906A0C8587" authorityName="Aspock &amp; Aspock" authorityYear="1991" box="[433,556,923,949]" class="Insecta" family="Berothidae" genus="Isoscelipteron" kingdom="Animalia" order="Neuroptera" pageId="6" pageNumber="137" phylum="Arthropoda" rank="species" species="borneole">
<emphasis id="77CCEAF7B003FFA0F5C76D906A0C8587" box="[433,556,923,949]" italics="true" pageId="6" pageNumber="137">I. borneole</emphasis>
</taxonomicName>
, but unlike in
<taxonomicName id="82B84D66B003FFA0F6BC6D906B0C8587" authorityName="Costa" authorityYear="1863" box="[714,812,923,949]" class="Insecta" family="Berothidae" genus="Isoscelipteron" kingdom="Animalia" order="Neuroptera" pageId="6" pageNumber="137" phylum="Arthropoda" rank="species" species="fulvum">
<emphasis id="77CCEAF7B003FFA0F6BC6D906B0C8587" box="[714,812,923,949]" italics="true" pageId="6" pageNumber="137">I. fulvum</emphasis>
</taxonomicName>
(
<bibRefCitation id="21294B14B003FFA0F74C6D976C698587" author="Aspock, U. &amp; Aspock, H." box="[826,1097,923,949]" pageId="6" pageNumber="137" pagination="65 - 76" refId="ref4196" refString="Aspock, U. &amp; Aspock, H. (1991) Zur Kenntnis des Genus Isoscelipteron Costa, 1863 (Berothidae: Berothinae). Zeitschrift der Arbeitsgemeinschaft Osterreich Entomologen, 43, 65 - 76." type="journal article" year="1991">Aspöck &amp; Aspöck 1991</bibRefCitation>
: fig. 14). We propose to assign the fossil into a new species in
<taxonomicName id="82B84D66B003FFA0F5826DB36AB285EB" authorityName="Costa" authorityYear="1863" box="[500,658,959,985]" class="Insecta" family="Berothidae" genus="Isoscelipteron" kingdom="Animalia" order="Neuroptera" pageId="6" pageNumber="137" phylum="Arthropoda" rank="genus">
<emphasis id="77CCEAF7B003FFA0F5826DB36AB285EB" box="[500,658,959,985]" italics="true" pageId="6" pageNumber="137">Isoscelipteron</emphasis>
</taxonomicName>
.
</paragraph>
</subSubSection>
</treatment>
</document>