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<mods:title id="249664519EA138FC43D63232DAA47818">A revision of the late Eocene snakeflies (Raphidioptera) of the Florissant Formation, Colorado, with special reference to the wing venation of the Raphidiomorpha</mods:title>
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<mods:namePart id="710B5C3FC76A457B2FF056342F02360C">Makarkin, Vladimir N.</mods:namePart>
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<mods:namePart id="4045C141580FB9D0BFBFA562AFD170B6">Archibald, S. Bruce</mods:namePart>
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<treatment id="039287A3FE0E371B23C76E92FF3BFEBD" ID-DOI="http://doi.org/10.5281/zenodo.6131141" ID-GBIF-Taxon="119420851" ID-Zenodo-Dep="6131141" LSID="urn:lsid:plazi:treatment:039287A3FE0E371B23C76E92FF3BFEBD" httpUri="http://treatment.plazi.org/id/039287A3FE0E371B23C76E92FF3BFEBD" lastPageId="9" lastPageNumber="410" pageId="8" pageNumber="409">
<subSubSection id="C321653EFE0E371A23C76E92FB5AF9F1" pageId="8" pageNumber="409" type="nomenclature">
<paragraph id="8B8436B5FE0E371A23C76E92FD6BFB44" blockId="8.[151,651,1196,1222]" box="[151,651,1196,1222]" pageId="8" pageNumber="409">
<heading id="D0CC81D9FE0E371A23C76E92FD6BFB44" bold="true" box="[151,651,1196,1222]" fontSize="11" level="1" pageId="8" pageNumber="409" reason="1">
<emphasis id="B94FEAA7FE0E371A23C76E92FD6BFB44" bold="true" box="[151,651,1196,1222]" pageId="8" pageNumber="409">
Family
<taxonomicName id="4C3B4D36FE0E371A23A66E92FD6BFB44" ID-CoL="6ZR" authority="Martynova, 1961" authorityName="Martynova" authorityYear="1961" box="[246,651,1196,1222]" class="Insecta" family="Baissopteridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Raphidioptera" pageId="8" pageNumber="409" phylum="Arthropoda" rank="family">
Baissopteridae
<bibRefCitation id="EFAA4B44FE0E371A22E56E92FD6BFB44" author="Martynova" box="[437,651,1196,1222]" pageId="8" pageNumber="409" refString="Martynova, O. M. (1961) Recent and fossil snakeflies (Insecta, Raphidioptera). Paleontologicheskii Zhurnal, 1961 (3), 73 - 83. [in Russian]" type="journal article" year="1961">Martynova, 1961</bibRefCitation>
</taxonomicName>
</emphasis>
</heading>
</paragraph>
<paragraph id="8B8436B5FE0E371A23C76ECCFE16FAD1" blockId="8.[151,1436,1266,2011]" pageId="8" pageNumber="409">
<emphasis id="B94FEAA7FE0E371A23C76ECCFD94FA89" bold="true" box="[151,628,1266,1291]" pageId="8" pageNumber="409">Diagnostic character states of venation.</emphasis>
Pterostigma usually with crossvein closing it proximally, with at least one incorporated branch of RA. Forewing: rich venation with 35 branches of
<collectingCountry id="F32C7625FE0E371A27426F29FBD4FAAC" box="[1042,1076,1303,1326]" name="Poland" pageId="8" pageNumber="409">RP</collectingCountry>
, many crossveins: 35 ra-pr, 5 15 ir, 47 r-m, 35 im, 23 icu.
</paragraph>
<paragraph id="8B8436B5FE0E371A23976F60FC5FFAF4" blockId="8.[151,1436,1266,2011]" box="[199,959,1374,1399]" pageId="8" pageNumber="409">
<collectingCountry id="F32C7625FE0E371A23976F60FF1FFAF5" box="[199,255,1374,1399]" name="India" pageId="8" pageNumber="409">Hind</collectingCountry>
wing: many crossveins: 45 ra-pr, 514 ir, 46 r-m, 24 m-cu.
</paragraph>
<paragraph id="8B8436B5FE0E371A23976FBCFB5AF9F1" blockId="8.[151,1436,1266,2011]" pageId="8" pageNumber="409">
<emphasis id="B94FEAA7FE0E371A23976FBCFEBFFA19" bold="true" box="[199,351,1410,1435]" pageId="8" pageNumber="409">Composition</emphasis>
. Five genera from the Cretaceous and Eocene:
<taxonomicName id="4C3B4D36FE0E371A20236FBCFB2DFA19" authority="Engel, 2002" authorityName="Engel" authorityYear="2002" box="[883,1229,1410,1435]" class="Insecta" family="Baissopteridae" genus="Cretoraphidiopsis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Raphidioptera" pageId="8" pageNumber="409" phylum="Arthropoda" rank="genus">
<emphasis id="B94FEAA7FE0E371A20236FBCFBDEFA19" box="[883,1086,1410,1435]" italics="true" pageId="8" pageNumber="409">Cretoraphidiopsis</emphasis>
<bibRefCitation id="EFAA4B44FE0E371A27156FBCFB2DFA19" author="Engel" box="[1093,1229,1410,1435]" pageId="8" pageNumber="409" refString="Engel, M. S. (2002) The smallest snakefly (Raphidioptera: Mesoraphidiidae): a new species in Cretaceous amber from Myanmar, with a catalog of fossil snakeflies. American Museum Novitates, 3363, 1 - 22. http: // dx. doi. org / 10.1206 / 0003 - 0082 (2002) 363 &lt;0001: tssrma&gt; 2.0. co; 2" type="journal article" year="2002">Engel, 2002</bibRefCitation>
</taxonomicName>
(monotypic) from Hauterivian/Aptian of Bon-Tsagaan,
<collectingCountry id="F32C7625FE0E371A21666F98FD47FA3D" box="[566,679,1446,1471]" name="Mongolia" pageId="8" pageNumber="409">Mongolia</collectingCountry>
;
<taxonomicName id="4C3B4D36FE0E371A21E26F98FC5DFA3D" authority="Willmann, 1994" authorityName="Willmann" authorityYear="1994" box="[690,957,1446,1471]" class="Insecta" family="Baissopteridae" genus="Lugala" higherTaxonomySource="GBIF" kingdom="Animalia" order="Raphidioptera" pageId="8" pageNumber="409" phylum="Arthropoda" rank="genus">
<emphasis id="B94FEAA7FE0E371A21E26F98FCE1FA3D" box="[690,769,1446,1471]" italics="true" pageId="8" pageNumber="409">Lugala</emphasis>
<bibRefCitation id="EFAA4B44FE0E371A20586F98FC5DFA3D" author="Willmann" box="[776,957,1446,1471]" pageId="8" pageNumber="409" refString="Willmann, R. (1994) Raphidiodea aus dem Lias und die Phylogenie der Kamelhalsfliegen (Insecta: Holometabola). Palaontologische Zeitschrift, 68, 167 - 197. http: // dx. doi. org / 10.1007 / bf 02989439" type="journal article" year="1994">Willmann, 1994</bibRefCitation>
</taxonomicName>
(monotypic) from the Early Cretaceous of Bayan-Tsagaan,
<collectingCountry id="F32C7625FE0E371A22016FF4FE21FA61" box="[337,449,1482,1507]" name="Mongolia" pageId="8" pageNumber="409">Mongolia</collectingCountry>
;
<taxonomicName id="4C3B4D36FE0E371A229D6FF2FCFBFA61" authority="Martynova, 1961" authorityName="Martynova" authorityYear="1961" box="[461,795,1482,1507]" class="Insecta" family="Baissopteridae" genus="Baissoptera" higherTaxonomySource="GBIF" kingdom="Animalia" order="Raphidioptera" pageId="8" pageNumber="409" phylum="Arthropoda" rank="genus">
<emphasis id="B94FEAA7FE0E371A229D6FF2FDB3FA61" box="[461,595,1484,1507]" italics="true" pageId="8" pageNumber="409">Baissoptera</emphasis>
<bibRefCitation id="EFAA4B44FE0E371A210A6FF5FCFBFA61" author="Martynova" box="[602,795,1482,1507]" pageId="8" pageNumber="409" refString="Martynova, O. M. (1961) Recent and fossil snakeflies (Insecta, Raphidioptera). Paleontologicheskii Zhurnal, 1961 (3), 73 - 83. [in Russian]" type="journal article" year="1961">Martynova, 1961</bibRefCitation>
</taxonomicName>
(twelve species) from pre-Barremian/early Barremian of Baissa, Transbaikalian
<collectingCountry id="F32C7625FE0E371A22F56FD0FE13F985" box="[421,499,1518,1543]" name="Russia" pageId="8" pageNumber="409">Russia</collectingCountry>
; Barremian/Aptian of Huangbanjigou, Yixian Formation,
<collectingCountry id="F32C7625FE0E371A27CF6FD0FB05F985" box="[1183,1253,1518,1543]" name="China" pageId="8" pageNumber="409">China</collectingCountry>
; late Albian of Spanish amber (Peñacerrada I); late Aptian of the Crato Formation of
<collectingCountry id="F32C7625FE0E371A20926C2CFBEAF9A9" box="[962,1034,1554,1579]" name="Brazil" pageId="8" pageNumber="409">Brazil</collectingCountry>
;
<taxonomicName id="4C3B4D36FE0E371A27486C2CFB57F9A9" box="[1048,1207,1554,1579]" class="Insecta" family="Baissopteridae" genus="Cretoraphidia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Raphidioptera" pageId="8" pageNumber="409" phylum="Arthropoda" rank="genus">
<emphasis id="B94FEAA7FE0E371A27486C2CFB57F9A9" box="[1048,1207,1554,1579]" italics="true" pageId="8" pageNumber="409">Cretoraphidia</emphasis>
</taxonomicName>
(four species) from Baissa and Neocomian of Romanovka, Transbaikalian
<collectingCountry id="F32C7625FE0E371A20576C08FCB5F9CD" box="[775,853,1590,1615]" name="Russia" pageId="8" pageNumber="409">Russia</collectingCountry>
;
<taxonomicName id="4C3B4D36FE0E371A20326C08FB2AF9CD" authority="Willmann, 1994" authorityName="Willmann" authorityYear="1994" box="[866,1226,1590,1615]" class="Insecta" family="Baissopteridae" genus="Austroraphidia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Raphidioptera" pageId="8" pageNumber="409" phylum="Arthropoda" rank="genus">
<emphasis id="B94FEAA7FE0E371A20326C08FBECF9CD" box="[866,1036,1590,1615]" italics="true" pageId="8" pageNumber="409">Austroraphidia</emphasis>
<bibRefCitation id="EFAA4B44FE0E371A27446C08FB2AF9CD" author="Willmann" box="[1044,1226,1590,1615]" pageId="8" pageNumber="409" refString="Willmann, R. (1994) Raphidiodea aus dem Lias und die Phylogenie der Kamelhalsfliegen (Insecta: Holometabola). Palaontologische Zeitschrift, 68, 167 - 197. http: // dx. doi. org / 10.1007 / bf 02989439" type="journal article" year="1994">Willmann, 1994</bibRefCitation>
</taxonomicName>
(monotypic) from the Crato Formation of
<collectingCountry id="F32C7625FE0E371A22CF6C64FE07F9F1" box="[415,487,1626,1651]" name="Brazil" pageId="8" pageNumber="409">Brazil</collectingCountry>
;
<taxonomicName id="4C3B4D36FE0E371A22A26C64FD7DF9F1" box="[498,669,1626,1651]" class="Insecta" family="Raphidiidae" genus="Dictyoraphidia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Raphidioptera" pageId="8" pageNumber="409" phylum="Arthropoda" rank="genus">
<emphasis id="B94FEAA7FE0E371A22A26C64FD7DF9F1" box="[498,669,1626,1651]" italics="true" pageId="8" pageNumber="409">Dictyoraphidia</emphasis>
</taxonomicName>
(monotypic) from the late Eocene of Florissant.
</paragraph>
</subSubSection>
<subSubSection id="C321653EFE0E371B23976C40FF3BFEBD" lastPageId="9" lastPageNumber="410" pageId="8" pageNumber="409" type="discussion">
<paragraph id="8B8436B5FE0E371A23976C40FD6AF8C9" blockId="8.[151,1436,1266,2011]" pageId="8" pageNumber="409">
<emphasis id="B94FEAA7FE0E371A23976C40FDF1F915" bold="true" box="[199,529,1662,1687]" pageId="8" pageNumber="409">Comments on characters.</emphasis>
The venation of
<taxonomicName id="4C3B4D36FE0E371A20516C40FC46F915" box="[769,934,1662,1687]" class="Insecta" family="Baissopteridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Raphidioptera" pageId="8" pageNumber="409" phylum="Arthropoda" rank="family">Baissopteridae</taxonomicName>
is in general similar to that of those
<taxonomicName id="4C3B4D36FE0E371A23C76C9CFEB3F939" box="[151,339,1698,1723]" class="Insecta" family="Mesoraphidiidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Raphidioptera" pageId="8" pageNumber="409" phylum="Arthropoda" rank="family">Mesoraphidiidae</taxonomicName>
(s.l.) that have a similar structure of the pterostigma (e.g., the Cretaceous genus
<taxonomicName id="4C3B4D36FE0E371A267F6C9CFE60F95D" authority="Ponomarenko, 1993" authorityName="Ponomarenko" authorityYear="1993" class="Insecta" family="Mesoraphidiidae" genus="Siboptera" higherTaxonomySource="GBIF" kingdom="Animalia" order="Raphidioptera" pageId="8" pageNumber="409" phylum="Arthropoda" rank="genus">
<emphasis id="B94FEAA7FE0E371A267F6C9CFA7CF939" box="[1327,1436,1698,1723]" italics="true" pageId="8" pageNumber="409">Siboptera</emphasis>
<bibRefCitation id="EFAA4B44FE0E371A23C76CF8FE60F95D" author="Ponomarenko" box="[151,384,1734,1759]" pageId="8" pageNumber="409" refString="Ponomarenko, A. G. (1993) Lower Cretaceous snakeflies from Transbaikalia. In: Ponomarenko, A. G. (Ed.), Mesozoic insects and ostracods from Asia. Trudy Paleontologicheskogo Instituta, 252, pp. 60 - 80. [in Russian]" type="journal article" year="1993">Ponomarenko, 1993</bibRefCitation>
</taxonomicName>
); it differs from these in possessing more branches of longitudinal veins and an enriched crossvenation. Therefore, the differences between this and other families are mainly numerical. The enriched venation of
<taxonomicName id="4C3B4D36FE0E371A22606D30FE35F8A5" box="[304,469,1806,1831]" class="Insecta" family="Baissopteridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Raphidioptera" pageId="8" pageNumber="409" phylum="Arthropoda" rank="family">Baissopteridae</taxonomicName>
is certainly secondary; Jurassic taxa of Raphidiomorpha (Metaraphidiidae and
<taxonomicName id="4C3B4D36FE0E371A23C76D0CFEB3F8C9" box="[151,339,1842,1867]" class="Insecta" family="Mesoraphidiidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Raphidioptera" pageId="8" pageNumber="409" phylum="Arthropoda" rank="family">Mesoraphidiidae</taxonomicName>
s.l.) have simpler venation.
</paragraph>
<paragraph id="8B8436B5FE0E371A23976D68FB46F858" blockId="8.[151,1436,1266,2011]" pageId="8" pageNumber="409">
<emphasis id="B94FEAA7FE0E371A23976D68FEDBF8ED" bold="true" box="[199,315,1878,1903]" pageId="8" pageNumber="409">Remarks.</emphasis>
We base our comments on examination of a single (Cretaceous) specimen of a species assigned to this family and published descriptions and figures of the others. We agree with
<bibRefCitation id="EFAA4B44FE0E371A27106D44FF00F835" author="Bechly" pageId="8" pageNumber="409" refString="Bechly, G. &amp; Wolf-Schwenninger, K. (2011) A new fossil genus and species of snakefly (Raphidioptera: Mesoraphidiidae) from Lower Cretaceous Lebanese amber, with a discussion of snakefly phylogeny and fossil history. Insect Systematics &amp; Evolution, 42, 221 - 236. http: // dx. doi. org / 10.1163 / 187631211 x 568164" type="journal article" year="2011">Bechly &amp; Wolf-Schwenninger (2011)</bibRefCitation>
that some of these published descriptions and figures appear problematic, and therefore our conclusions (diagnosis and family composition) are preliminary pending extensive revision of these taxa.
</paragraph>
<paragraph id="8B8436B5FE0F371B23976AA9FF3BFEBD" blockId="9.[151,1436,151,319]" pageId="9" pageNumber="410">
The family is sometimes considered to be paraphyletic (e.g.,
<bibRefCitation id="EFAA4B44FE0F371B20D46AA9FBD6FF32" author="Willmann" box="[900,1078,151,176]" pageId="9" pageNumber="410" refString="Willmann, R. (1994) Raphidiodea aus dem Lias und die Phylogenie der Kamelhalsfliegen (Insecta: Holometabola). Palaontologische Zeitschrift, 68, 167 - 197. http: // dx. doi. org / 10.1007 / bf 02989439" type="journal article" year="1994">Willmann 1994</bibRefCitation>
;
<bibRefCitation id="EFAA4B44FE0F371B27146AA9FF32FF56" author="Bechly" pageId="9" pageNumber="410" refString="Bechly, G. &amp; Wolf-Schwenninger, K. (2011) A new fossil genus and species of snakefly (Raphidioptera: Mesoraphidiidae) from Lower Cretaceous Lebanese amber, with a discussion of snakefly phylogeny and fossil history. Insect Systematics &amp; Evolution, 42, 221 - 236. http: // dx. doi. org / 10.1163 / 187631211 x 568164" type="journal article" year="2011">Bechly &amp; Wolf-Schwenninger 2011</bibRefCitation>
), as it is not defined by any distinct synapomorphies. We did not find these as well. It is unknown yet if this group of genera with enriched venation constitutes a monophyletic taxon or not, perhaps within a paraphyletic
<taxonomicName id="4C3B4D36FE0F371B23C76B3AFEB3FE9F" box="[151,339,260,285]" class="Insecta" family="Mesoraphidiidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Raphidioptera" pageId="9" pageNumber="410" phylum="Arthropoda" rank="family">Mesoraphidiidae</taxonomicName>
(s.l.). Both families are in strong need of detailed revision, but this is beyond of the scope of this paper.
</paragraph>
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